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PROCEEDINGS 


of the 


Biological Society of 


Washington 


VOLUME 96 


1983 
Vol. 96(1) published 25 March 1983 Vol. 96(2) published 7 July 1983 


Vol. 96(3) published 31 October 1983 Vol. 96(4) published 29 December 1983 


WASHINGTON 
PRINTED FOR THE SOCIETY 


EDITOR 


BRIAN KENSLEY 


ASSOCIATE EDITORS 


Classical Languages Invertebrates 

GEORGE C. STEYSKAL THOMAS E. BOWMAN 

Plants Vertebrates 

DAVID B. LELLINGER RICHARD P. VARI 
Insects 


ROBERT D. GORDON 


All correspondence should be addressed to the 
Biological Society of Washington, Smithsonian Institution 
Washington, D.C. 20560 


ALLEN PRESS INC. 
LAWRENCE, KANSAS 66044 


OFFICERS AND COUNCIL 
of the 
BIOLOGICAL SOCIETY OF WASHINGTON 
FOR 1982-1983 


OFFICERS 
President 
PAUL J. SPANGLER 


Vice President 
DAVID L. PAWSON 


Secretary 
CATHERINE J. KERBY 


Treasurer 
LESLIE W. KNAPP 


COUNCIL 


Elected Members 
JANET C. GOMON RICHARD S. HOUBRICK 
GORDON HENDLER ROY W. McDIARMID 
JAMES C. TYLER 


TABLE OF CONTENTS 


Volume 96 


Alvarino, Angeles. Nectocarmen antonioi, a new Prayinae, Calycophorae, Siphonophora 
Aire rns © ea a Te a Fa aes ee A coe lace elas Oe ec ee 
Alvarifio, Angeles, and Kenneth R. Frankwick. Heteropyramis alcala and Thalassophyes 
ferrarii, new species of Clausophyidae (Calycophorae: Siphonophorae) from the South 
ACHETER Te 2 SO ei ae AN ME AGEN ert NE A eee Re aC ee 
Bieri, Robert, Dolores Bonilla, and Fernando Arcos. Function of the teeth and vestibular 
organ in the Chaetognatha as indicated by scanning electron microscope and other 
ODSC EVAL OM Ser Se rare cs SI ae, ls) ei Pe Ste 
Bongers, Tom. Revision of the genus Leptosomatum Bastian, 1865 (Nematoda: 
Weptosomiatidae) 2s. ok Se leet ater Ng ee RS ee Ba ech a ee 
Bowman, Thomas E. III, and H. H. Hobbs, III. Caecidotea filicispeluncae, a new troglobitic 
asellid asopodtfrom:@hiozs) 202 ee ee ee, ee ee 
Bowman, Thomas E., and Thomas M. Illiffe. Bermudalana aruboides, a new genus and 
species of troglobitic isopoda (Cirolanidae) from marine caves on Bermuda... 
Brinkhurst, Ralph O., and R. Deedee Kathman. Varichaetadrilus, a new name for Vari- 
chaeta Brinkhurst, 1981, non Speiser, 1903, (Diptera) with a description of a new species 
Wa fPALT CID oN co I NS le NR SNe SRR A ce SOc a 
Brinkhurst, Ralph O., and R. Deedee Kathman. Arctodrilus wulikenses, new genus, new 
species (Oligochaeta: Tubificidae) from Alaska cece ccecceeeeceeeeeeeeeeeeeeetteetteeneens 
Bueno-Soria, Joaquin. Three new species of Ochrotrichia (Metrichia) from Chiapas, Mexico 
(Erichoptera) Hydroptilidae) ee ee 
Burr, Brooks M., and Walter W. Dimmick. Redescription of the Bigeye Shiner, Notropis 
boons (RISCES: Cyprinidae) es eee a ee 
Cairns, Stephen D. Observations of species of the fossil genus Axopora (Coelenterata: 
Hydrozoa) and its evolutionary significance to the Stylasteridae 
Calder, Dale R. Hydroida from estuaries of South Carolina, USA: Families Sertulariidae 
and -Plumvuslariidaé: 22.2605 22 385 oo A eS 
Cannatella, David C. Synonymy and distribution of Phyllomedusa boliviana Boulenger 
(Anura? Hylidae) 20.52 eee es ee ee 
Child, C. Allan. Pycnogonida of the western Pacific islands II. Guam and the Palau 
SVAN LS aeolian ea Ee oC ee 8 UR eh 
Coates, Kathryn A. A contribution to the taxonomy of the Enchytraeidae (Oligochaeta). 
Review of Stephensoniella, with new SPeCieS LeCOLGS nec ecccceeeeeeeecececeeeeeeeeeeeeeennneeeeeeeeeeeeennneeeeeeness 
Coen, Loren D., and Kenneth L. Heck, Jr. Notes on the biology of some seagrass-dwelling 
crustaceans (Stomatopoda and Decapoda) from Caribbean Panama 
Collette, Bruce B. Recognition of two species of double-lined mackerels (Grammatorcynus: 
Scomaby rae) pe) eee en ee te ee ne 
Collete, Bruce B. Two new species of coral toadfishes, family Batrachoididae, genus San- 
opus trompYaucatans Mexico amg) Belize ee 
Cressey, Roger F., and Marilyn Schotte. Three new species of Colobomatus (Copepoda: 
Philichthyidae) parasitic in the mandibular canals of haemulid fishes... 
Crumly, Charles R. An annotated checklist of the fossil tortoises of China and 
ING OLGA ee Sha ne a Dee ee ANN ie No ee 
Cutler, Edward B., Norma J. Cutler, and Peter E. Gibbs. A revision of the Golfingia 
subgenera Golfingiella, Stephen, 1964, and Siphonoides, Murina, 1967 (Sipuncula)........ 
Erséus, Christer. A new bathyal species of Coralliodrilus (Oligochaeta: Tubificidae) from 
the southeast: Atel cir tc os AEE Le aR eee oe 
Fauchald, Kristian. Life diagram patterns in benthic polychaetes... 
Felgenhauer, Bruce E., and Joel W. Martin. Atya abelei, a new atyid shrimp (Crustacea, 
Decapoda, Atyidae) from the Pacific slope of Pamarma occ eeeeeececceeceeneeeeeeeeeeeeeeeneeeeeeeee 
Fitzhugh, Kirk. New species of Fabriciola and Fabricia (Polychaeta: Sabellidae) from 


339-348 


686-692 


110-114 


807-855 


693-697 


291-300 


301-306 


407-410 


79-83 


50-58 


758-769 


7-28 


59-66 


698-714 


411-419 


202-224 


715-718 


719-724 


189-201 


567-580 


669-674 


273-275 
160-177 


333-338 


276-290 


Fleminger, A. Description and Phylogeny of Jsaacsicalanus paucisetus, n. gen., n. sp., 
(Copepoda: Calanoida: Spinocalanidae) from an east Pacific hydrothermal vent site 


Flint, Oliver S., Jr. Studies of neotropical caddisflies, XX XIV: the genus Plectromacronema 
@inichopterashy dropsychidac) “sss ee ee ee ee 
Franz, Richard, and Horton H. Hobbs, Jr. Procambarus (Ortmannicus) leitheuseri, new 
species, another troglobitic crayfish (Decapoda: Cambaridae) from peninsular Florida.. 
Gardner, Alfred L. Proechimys semispinosus (Rodentia: Echimyidae): Distribution, type 
locality sand taxonomic isto rye Oe eee eS eet he Beer apel ene wks 
Gore, Robert H. The identity of Petrolisthes marginatus Stimpson, 1859, and the descrip- 
tion of Petrolisthes dissimulatus, n. sp. (Crustacea: Decapoda: Porcellanidae).................... 
Grant, Richard E. Argyrotheca arguta, a new species of brachiopod from the Marshall 
KSlandssswestennieacihi crvevsy tamer a bees able ele BO nel eee TN Pel Won eno Be, 
Heaney, Lawrence R., and Robert M. Timm. Systematics and distribution of shrews of 
the genus Crocidura (Mammalia: Insectivora) 1m Vietrra rm cic cece ee ceecceeeneeees 
Heemstra, Phillip C., and William D. Anderson, Jr. A new species of serranid fish genus 
Plectranthias (Pisces: Perciformes) from the southeastern Pacific Ocean, with comments 
GUN CINUS WE CLICIOIO Sh pers Get aes ie eo OR Been inr Aeecvy Tew SL decries oo 
Hendrickx, M. E., M. K. Wicksten, and A. M. van der Heiden. Studies of the coastal 
marine fauna of southern Sinaloa, Mexico. IV. Report on the caridean crustaceans ........ 
Heyer, W. Ronald. Clarification of the names Rana mystacea Spix, 1824, Leptodactylus 
amazonicus Heyer, 1978 and a description of a new species, Leptodactylus spixi (Am- 
phibiayweptodactylidae)=7 00) £On t Wels cole ee ee eet ee 
Heyer, W. Ronald. Notes on the frog genus Cycloramphus (Amphibia: Leptodactylidae), 
with descriptions Of tWO MEW SPEECHES ..nna.ecccccccccceececccccceccseseececcessssssssesecsesssstsessseeeesesssonessessesssssteeesseseeesestensseeesseseee 
Heyer, W. Ronald, and William F. Pyburn. Leptodactylus riveroi, a new frog species from 
Amazonia, South America (Anura: Leptodactylidae) 2.0 
Hobbs, Horton H.., Jr. Distocambarus (Fitzcambarus) carlsoni, anew subgenus and species 
of crayfish (Decapoda: Cambaridae) from South Carohima i... ccccccceccecccccceccccececeeeeeeeeeeeseeeeee 
Hobbs, Horton H., Jr., and Paul H. Carlson. Distocambarus Decapoda: Cambaridae) 
elevated to generic rank, with an account of D. crockeri, new species, from South 
2 lina cree ee ee ee Ee ea ek a mene Pee cnet Eisele 
Hobbs, Horton H., Jr., and Auden C. McClure. On a small collection of entocytherid 
ostracods with the descriptions of three mew SPeCLeS eee 
Houbrick, Richard S. A new Strombina species (Gastropoda: Prosobranchia) from the 
cROpicalawestenn “Atlan iG... Ae el SU uu a ete Ley Satay Bote ik er ep Dl uke 
Huddleston, Richard W., and Kurt M. Savoie. Teleostean otoliths from the late Cretaceous 
(Maestrichtian age) Severn formation of Maryland 
Jewett, Susan L., and Emest A. Lachner. Seven new species of the Indo-Pacific genus 
ERVIOLG(RISCESS GObLIGaAC) Maree Amami ost Pekin feb be SN Bein eck SN ee 
Kornfield, Irv, and Jeffrey N. Taylor. A new species of polymorphic fish, Cichlasoma 
minckleyi, from Cuatro Ciénegas, Mexico (Teleostei: Cichlidae) 0 
Kornicker, Louis S. Harbansus slatteryi, a new species of myodocopine ostracode from 
the Great Barrier Reef of Australia (Plilormedidace) 2.00... cccccccecccccceeeeeeeecccnsneeeeeeeeeeventeeeeeeeeee 
Kornicker, Louis S. Zeugophilomedes, a new genus of myodocopine ostracode 
(Rhilomredinac) seis r thes bceley este AEN Dd SSE eet lene eae DY yl eaakiayly to Goel it 
Kritsky, Delane C., and Vernon E. Thatcher. Neotropical Monogenea. 5. Five new species 
from the Aruana, Osteoglossum bicirrosum Vandelli, a freshwater teleost from Brazil, 
with the proposal of Gonoclethrum n. gen. (Dactylogyridae: Ancyrocephalinae)...... 
Kudenov, Jerry D. Streptospinigera heteroseta, a new genus and species of Eusyllinae 
(Polychaeta: Syllidae) from the western shelf of Florida 0.000000. 
Lapota, David. Bioluminescence in the marine ostracod Cypridina americana (Miiller, 
1890) off Manzanillo, Mexico (Myodocopa: Cypridiminae) 2.2... 
Lewis, Julian J. The assignment of the Texas troglobitic water slater Caecidotea pilus to 
the genus Lirceolus, with an emended diagnosis of the genus (Crustacea: Isopoda: 
PARSE Uf GUE G) ee RU ee GER eee te acre Mah a ha ti De ad We el che AN et eth WT ek aes 
Lewis, Julian J. Caecidotea fonticulus, the first troglobitic asellid from the Ouachita Moun- 
tains i(@rustaceas:Isopoda>Asellidae))..05) ee 


605-622 


225-237 


323-332 


134-144 


89-102 


178-180 


115-120 


632-637 


67-78 


270-272 


548-559 


560-566 


429-439 


420-428 


770-779 


349-354 


658-663 


780-806 


253-269 


181-188 


478-480 


581-597 


84-88 


307-308 


145-148 


149-153 


Manning, Raymond B., and David K. Camp. Fennerosquilla, a new genus of stomatopod 
crustacean from the northwestern Atlantic 2. cece ceeeeeeeeeecneeeneneenmnnnceneeenenennunnne 
Markle, Douglas F., Jeffery T. Williams, and John E. Olney. Description of a new species 
of Echiodon (Teleostei: Carapidae) from Antarctic and adjacent seas 
McKenzie, K. G. Bonaducecytheridae McKenzie, 1977: A subjective synonym of Psam- 
mocytheridae Klie, 1938 (Ostracoda: Podocopida: Cytheracea) 0.00. 
Nelson, Joseph S. Creedia alleni and Creedia partimsquamigera (Perciformes: Creediidae), 
two new marine fish species from Australia, with notes on other Australian creediids... 
Olson, Storrs L. Geographic variation in Chlorospingus ophthalmicus in Colombia and 
Venezuelay(Aves= Thraupidac) i. sa ere. Pa Be ee el eee ee ee 
Pawson, David L. Psychronaetes hanseni, a new genus and species of elasipodan sea 
cucumber from the eastern central Pacific (Echinodermata: Holothuroidea)... 
Petit, Richard E. A new species of Cancellaria (Mollusca: Cancellariidae) from the northern 
Gulf Of MI@xi COM te ts lS 2 Bees als) 2 Pee SA MLE TD od ct eee ta a 
Pettibone, Marian H. A new scale worm (Polychaeta: Polynoidae) from the hydrothermal 
TGLif t= LTS AMO Alma SS Le Tem NVA © Kak Gretta NI Pacem unc nNOS ESE 
Pettibone, Marian H. Minusculisquama hughesi, a new genus and species of scale worm 
(Polychaeta: Polynoidae) from eastern Camada cece 
Poss, Stuart G., and Victor G. Springer. Eschmeyer nexus, a new genus and species of 
SCORPASITA AST may EY he UE See re Ce ge Ee 
Quinn, James F., Jr. Carenzia, a new genus of Seguenziacea (Gastropoda: Prosobranchia) 
WithEthe GeSGuip tlOnKo lease WAS [CCI eC Seema cee et nnn us enn eS 
Quinn, James F., Jr. A revision of the Seguenziacea Verrill, 1884 (Gastropoda: Proso- 
branchia). I. Summary and evaluation of the superfamily 
Sakai, Tune. Eight new species of Indo-Pacific crabs from the collections of the Smithsonian 
ys tin uns Ta ES gD IAL 0 OE aaa tee, COE ee 
Schultz, George A. Disposition of three species of Oniscoidea from western Atlantic sea- 
shores (Crustacea: Isopoda: Halophilosciidae and Philosciidae) 0... 
Schultz, George A. Two species of 7y/os Audouin from Chile, with notes on species of 
Tylos with three flagellar articles (Isopoda: Oniscoidea: Tylidae) 
Shelley, Rowland M. Parvulodesmus prolixogonus, a new genus and species of xystodesmid 
milliped from South Carolina (Polydesrmica) 000. eeeeeeeeennnennnneeeteeeeeeeeeneeenee 
Solis-Weiss, Vivianne. Parandalia bennei (Pilargidae) and Spiophanes Iowai (Spionidae), 
new species of polychaetous annelids from Mazatlan Bay, Pacific coast of Mexico... 
Thomas, James Darwin. Curidia debrogania, a new genus and species of amphipod (Crus- 
tacea: Ochlesidae) from the barrier reefs of Belize, Central America... 
Toll, Ronald B. The lycoteuthid genus Oregoniateuthis Voss, a synonym of Lycoteuthis 
Pfeffer, 1900 (Cephalopoda: Teuthoidea) ccc ecceccececeeceeceeeeeeeeeeeteeeesttetipnnttnnnnnnnneeneeeeene 
Webster, J. Dan. A new subspecies of fox sparrow from Alaska 
Weems, Robert E., and Stephen R. Horman. Teleost fish remains (Osteoglossidae, Blo- 
chiidae, Scombridae, Triodontidae, Diodontidae) from the Lower Eocene Nanjemoy 
hormationsof Maryland)...wso8 eee ee a ee ee eee 
Wicksten, Mary K., and Terrance H. Butler. Description of Evalus lineatus, new species, 
with a redescription of Heptacarpus herdmani (Walker) (Caridea: Hippolytidae) ............ 
Williams, Austin B., and Cindy Lee Van Dover. A new species of Munidopsis from 
submarine thermal vents of the East Pacific Rise at 21°N (Anomura: Galatheidae)... 
Williams, J. T., and A. M. Smart. Redescription of the Brazilian labrisomid fish Starksia 
brasili€nsis:se Seek Meek, Sher years, Se bo 2 LAN EOD yt A CoN ells ea ie SUN RR, 
Williams, N. Sandra. Three new species of Brueelia (Mallophaga: Philopteridae) from the 
Miumiudae(Awes: Passeriformes): met ss oe ie el Se 
Wilson, George D. F. An unusual species complex in the genus Eurycope (Crustacea: 
Isopoda: Asellota) from the deep North Atlantic Ocean... ccccecccceccceceeeeeeeeeeeececettneeeeeeeneeeeese 
Wirth, Willis W., and William L. Grogan, Jr. The Nearctic species of the Bezzia bivittata 
group (Diptera: Ceratopogomidae). i ceccccceeecccccsccccceseeeeeeeeeeesseeeesesevitesutuncnceceseeeeeseeseesestenesssustessesuuete 
Wolf, Paul S. A revision of the Bogueidae Hartman and Fauchald, 1971, and its reduction 
to Boguelinae, a subfamily of Maldanidae (Polychaeta) 2000.00.00... cccccscccsssscsecceeeeeeessesseessesssteeennenee 


317-322 


645-657 


684-685 


29-37 


103-109 


154-159 


250-252 


392-399 


400-406 


309-316 


355-364 


725-757 


623-631 


440-451 


675-683 


121-126 


370-378 


127-133 


365-369 


664-668 


38-49 


1-6 


481-488 


638-644 


599-604 


452-467 


489-523 


238-249 


Worley, E. K., and David R. Franz. A comparative study of selected skeletal structures 

in the seastars Asterias forbesi (Dosor), A. vulgaris Verrill, and A. rubens L., with a 

discussion of possible relatiOrSb ps ee cece eeeeeeeeeeeceeeeeeeeeeeecennnnnnnnnnnetnneeeneneeeennnneeeee 524-547 
Zottoli, Robert. Amphisamythia galapagensis, a new species of ampharetid polychaete 

from the vicinity of abyssal hydrothermal vents of the Galapagos Rift, and the role of 

CIMISESPECIES MMnIt tECOSY SLETING eta ss nee ee ed es 379-391 
Zullo, Victor A., and Jon D. Standing. Sponge-inhabiting barnacles (Cirripedia: Archaeo- 

balanidae) of the Carolinian Province, southeastern United States, with the description 

of a new species of Membranobalanus Paspry i... eee ennnnnnnnnncnnsesnseceeeeeeeeeeeeeeee 468-477 


INDEX TO NEW TAXA 


VOLUME 96 


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


COELENTERATA 
Axoporarcleith mi dauiiadiic: ue sees 2 ee ed ee ee 764 
Heteropyramis Q/cala ee 686 
Nectocarmen 339 
TUL OTTO VR hat S catego ng SE ees ota tee ene Oe ore eer aR OA oe 339 
‘hhalassophy es ferGril.s-24.\ 27.28 eh SS ees ee a) eee 689 
‘iPhyroscyphusmmareinatus NiCr) 2 tee es ee ee 16 
PLAT YHELMINTHES 
Trematoda 

(GOVOCIOU EI UTY Nee ees a Fan eer al Su Re eee ee cee 586 
CONTA LUO 10 cS ee SR CR Bs nT ag EY ee Ne APN UIE 587 
COCTIOVA CUTTERS #0 os esa ee aes a ori a a eae. et ees ee 590 

CULTIST LIAS eco a cc er erence pe shrsrcte ae thn orange te nrc eda 592 
DIGTGCT OUD CUTTY SE re Ne ee 592 

DP LUTAGLGT USS 2 tte? 0 persia Sea OG Oe tao eee Ne ea OL ae i en le 587 

NEMATODA 
Leptosomatumi isundacnse mn: Cs ok eee eee Steere 844 
ANNELIDA 
Polychaeta 

Amphisamythavealapagensis 22) ee id ee 382 
air Glatt flO GUGUGs eho ein MY nel ld a ole i Sr 284 
Fabriciola trilobata 276 
Lepidonotopodiumn....... 392 
fimbriatum......... 393 
VAGUE GUILDS GULQTIUG ci eal sh UW eh RUIN ret, Ss A eM et cect ele 400 
PLUG TV es (etmek ETL a ee 401 
Parandaliawbennetis xo. 2 2s: cue Re oe ee 2 ac Re ee dale 370 
Spropmanesw/OWai coos eae el all pe yd dae site ade een res Oe 373 
Streptospinigera 84 
WC CV OS CL ee Nee rece sa EE a BE EN eel Soa alee eee ant ee 85 

407 

408 

CoOralliodrilus Lovicic ict vss este st oa tae el ee 273 
Stephensoniellai Sterne terse: koe Ss ik lane els noe eal ee Oe ae he 415 
UTS Vi OTM ATM Cece a Ns IR Ba at Ya lt od oe ad 416 
Varichaétadrilus (ipl epi sccten A Ws ae Oe arpa ade ta a 2 es 302 


ATPYTOUNCCA Greta 2 Ne 2 Sm eee ee Das 52 Gs les, gee se 8 


ARTHROPODA 


Crustacea 


PSNI AME ED C1 Cotesia Rr er SOc Ue ree eR tha re Ae neon ericrtern nO Net ares Auveel sulted Oy Ne Bale 
Ankylocythere carpenteri 
Ascetocythere jezerinaci......... 

IB CHPREADUAY LICH ICH AYGT Nes ok oe en NPC TT Ae gROR a ete Noe EAN Ns AO A ae Se nee DT 
AT UIDOTEL CS eee er eerie seer ae tet Rie abe aed i eee Te a Ne an Ne ys i ly 

Caecidotea filicispeluncae 
fonticulus......... 

Cancer /uzonensis................. 

Col oboniatusHp eli Zeris iS pee ee erecta eee ase ten er hn ce Deke a ee Ee eee 
(GCL D0) Cea etek TN aetna ROI Re So eR ed Oa ad un Des Sh leal eee ea OI 
LLL IT CL Tg1 LS Pac eae OPA ROR ISSR OER a EM aso ae Me wR tA Ce ee eee 

GinPLOCMEIMUSHARCINALUS ass ae ee lh 

Curidia 
ODDO IANS aoe I EI SS aa, SF DePUL OA Nt TT cto ee WOT OS 

Distocamibanus (Hitzcambarus) 22 he ee Ue oe 
CNIS OUTO ss SSA SS les a aN UPA A, SE Silt en aU ODT EU ete 
(CT OG ICE TJ ten aren Ee NN Ele Nf nd oN aE a A i inte ole oa ee Nati 

Ebalia punctulata............... 

Ethusa philippinensis 

Eualus Jineatus ................... 

ESUEGVICODICNCH ECU LETIC CL ae rte ORak nian tiie ag Mea tical SNS Ace MRT A Fy deeper JS eas nee ER 
BLS CE amenemed Si lai a oka Retro pee pe ROE SRD eae etn WE Pra Res Slade secas ND tinh oa ee Mea A 
LESS (C1 ieee aan aes cee ae cies ty ee A PUL on se Se Ey ld ae aint Pe Dee ta cede as 
[DESCRIBES SOP SI ec RIM Desa Fee Pe 4 ALN rd DoCS or oO 

Fennerosquilla 

VAT OAH SUSHS/ LLC Ig) Leese re AUN PS Mare 0) a I UR On Reece OO Lu (Nea We Le eho 

SUIS CEES UGH CULES Peseta ete te AMIEL EN Sa IC a a oa tele a ater ey ee rs al ele Mee ALUN 
PPD EULCTS CLUS ers ae ee ee eee RL Ned SN AS hl AS ee So ere g Us OSA NS RS late IER Oe PN 

NECUGCOSIAM ITIL CES U Sento es OVC ree el UA aN ol a laeemvbasrat oS aes Otte Seek Taka Un eRe 

Membranobalanus costatus 

INMtnT Gl OPSISR/ CTL ISO mares reese Alen ee a Wh eal el es ese ey nt Ga a PR Cone 

TMC O Gy EST HLTH Ne NG RI wl aca eee lh oss taeda) SN Ee ee eae 

AIC USHOLC TIT CELLS etme nittan cere eI AL NE YS ERB Aly ib nse enor Muro Tes NE Ne Gea 


Praebebalia septemspinOsa ccc 
Procambarus (Ortmannicus) /Jeitheuseri 
Pseudophilyra Dur Mensis. ii... 

BS CLO SCL CL aaemaaen tral NTP a aa At ANT NEF Mon SIN Pad ae eek oe koe al UN keen PPE WS 
BL SH 2115 15 sree aeRO rare od WA esol SC A 2 ol eri Aine enh Wa a A San er 
EUS OP ILO TILE AC SIPMSU SA soll AME NAD AMEE BR MLS PSU uta Ma 2a ANN x aso beh aoe YE ae 


ISG ZZN ANIC AVERCHIS 1 Steremee es ete NGS ie eee ns OE I I er ea ene 
andersonorum 
CLT UE CEE CET E i eee aly ato anne On a ce tay Ree ETE pe eM Ue ee etna ge 
CY QTISIS HUG esse RE NSIT ete URE NI ra sear Mea eer AU cd ON et ee ee aie PEE La DLC ABE CORA Oy 
gibberella........ 
luteiventris 
mohave ............ 

LUT UD CS pete ata Red be ree Nee ee ol Rete eae ne Ln Za Ae oP aL aoe the See 
sandersoni 
Setisinotumi......... 
spathula....... 
GNC TES TS reset ee nr eg De tac eae ee SRL UE ct 19 Te ch eh a gel te lc, atoll ea 


Brueelia dorsale 600 


montana............-- 600 
polyglotta 602 
Ochnotnichial (Wektichia)\@\0/= = ee ee 82 
lacuna 79 
TW GS so ene etch 79 
Plectromacronema /isae 228 
POV Ub OG OSTIUUIS ite hese 80 Nc Sea Ma eR AR SG dP he ere es Re 121 
DIV OUIX OS OMUS foes este decree ON ae teal 0 i Nae cn a ie 

Pycnogonida 
Arm othe ait ia po val cass le ae ee) ie loa meet lin ln 701 
Anoplodactylus; CHAMONTUS icc Se ea te ence 705 
AUIStrOGECUS PQLQUCTIS Core ons ee nee ee einen de a ek ee lal 711 
Sequapallene wicrone@sica@ :s.2.20.0 ceo ee ee tee Se ee 709 

MOLLUSCA 
GamCellariaytOSC Water sce) haces a al ao ak Ate as Leal en cae ee oe ca 250 
QT TZ he oe psa Na Sha nA nA ae CO a gern gi Se nore i 355 
TLC TUS eee oe Wa a Se de 362 
Strombina (Cotonopsis) argentea 349 
PS CHY ONGCEOS io) 8 lth dt A ee a ca ar pa eee lee 154 
PLTAS C1Ai i 0 oho i en aA ele Seles el 155 

CHORDATA 

Pisces 
GichlaSomia7i7Clel Cyt Sic eto Ot eI Be ue Aa 254 
Creedia Qe ni 2.0... 29 
partimsquamigera 33 
Echiodon cryomargarites 647 
SENS CHITVUCW CH ise a nF IE La hoy EN LL rs dR lesa le Ae Os Ae Se 309 
TLCS one ei, Bed On sso ae Sng ot cS) ta Sa ea ine et ole ee 310 
Ewiota al bolineata 20). eee Oe ee EE eee 783 
CONMMCL G20 oe ak at FI MR SO ae 796 
japonica 788 
VOR GQSCLALAE 28, 2 Nei ree ON Vice NS SE aR Ree AS CURE ieee: Sa, Ve OR a 791 
DUCE Gt Ae rk iee ot MS ES SO Ss 793 
SOU OT Gees UC Ba Be a tee eee ee eet Be ee 799 
ES TV LTS Cte a i Ae cece het lo Pal Ms ce ee reer a At ee ear 802 
Plectranthias exsul 632 
SanopustercenticldOrun 20.2 oak eee 721 
TELUGU ATUS teres Bec Es, iL rl Se Se en ee 719 
Amphibia 
Gycloramiphiisie@edrensis 2s te ek Nee ae ee 553 
izecksohni 550 
Leptodactylus riveroi 560 
SDD Uh ec lM Re et ne el EL. Noo GR 270 
Aves 

Chlorospingus ophthalmicus exitelus 3 108 
ERDAS oe 8 eae eee 107 


Passerella aliaca seh il Cat eris 1 Sess ee ee ri ale UT a 664 


(ISSN 0006-324) 


Proceedings 
| of the 
BIOLOGICAL SOCIETY 
ne ( 
WASHINGTON 


7 SMITASOR 
APR 4 sap- 
wird 983 
CBP 


ARIES 


Volume 96 25 March 1983 Number 1 


i 


THE BIOLOGICAL SOCIETY OF WASHINGTON 


1982-1983 
Officers 
President: Paul J. Spangler Secretary. Catherine J. Kerby 
Vice President: David L. Pawson Treasurer: Leslie W. Knapp 
Elected Council 
Janet C. Gomon Richard S. Houbrick 
Gordon Hendler Roy W. McDiarmid 


James C. Tyler 


Custodian of Publications: Michael J. Sweeney 


PROCEEDINGS 
Editor: Brian Kensley 


Associate Editors 


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PROC. BIOL. SOC. WASH. 
96(1), 1983, pp. 1-6 


DESCRIPTION OF EUALUS LINEATUS 
NEW SPECIES, WITH A REDESCRIPTION 
OF HEPTACARPUS HERDMANI (WALKER) 
(CARIDEA: HIPPOLYTIDAE) 


Mary K. Wicksten and Terrance H. Butler 


Abstract.—A small striped hippolytid shrimp has been identified as Eualus 
herdmani by several authors. Examination of the holotype of Spirontocaris herd- 
mani Walker revealed that this unique specimen should be assigned instead to 
the genus Heptacarpus, and that it differs from an undescribed species assigned 
to it in the literature. Heptacarpus herdmani is redescribed. The common subtidal 
striped shrimp of the western coast of North America is described herein as a 
new species, E. lineatus. 


Species of Eualus live in boreal to cool temperate waters of the northern and 
southern hemispheres (Wicksten 1979). A small striped shrimp of this genus has 
been collected subtidally along the Pacific coast from Alaska to western Mexico. 
This shrimp has been identified as E. herdmani (Walker). However, specimens 
of this animal do not agree completely with the description given by Walker 
(1898). We borrowed specimens from the Allan Hancock Foundation (AHF), and 
the U.S. National Museum (USNM) to compare them with the holotype, bor- 
rowed from the British Museum (Natural History). 

This paper describes as a new species E. lineatus, previously misidentified as 
E. herdmani. Spirontocaris herdmani is redescribed and assigned to Heptacar- 
pus. 

This work was supported in part by a fellowship from the Allan Hancock 
Foundation and a minigrant from Texas A&M University. Figures 1—2 were pre- 
pared by Alastair Denbigh, Pacific Biological Station. 


Heptacarpus herdmani (Walker) 


Spirontocaris herdmani Walker, 1898:277—278, pl. 16, fig. 2. 

Heptacarpus herdmani.—Holmes, 1900:204. 

Spirontocaris herdmani.—Rathbun, 1904:100-101 (in part). 

Eualus herdmani.—Holthuis, 1947:11 (in part)—Word and Charwat, 1976:103-— 
104 Gin part).—Butler, 1980:197—198 (in part). 

Not Eualus herdmani of Hobson and Chess, 1974:579 (=Eualus lineatus n. sp.) 


Description.—Rostrum horizontal, reaching beyond second segment of anten- 
nular peduncle and almost to end of scaphocerite. Two dorsal teeth on carapace, 
series of 3 to end of eyestalk, distal half of rostrum without dorsal teeth. Apex 
of rostrum sharp. Rostrum with | prominent ventral tooth just behind tip, slight 
knob behind tooth. 

Carapace smooth, dorsum straight. Very small knob at level of suborbital spine. 
No supraorbital spines. Antennal spines strong, prominent; pterygostomian spine 
very small and sharp. 


i) 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Abdomen smooth. Pleurae of segments 1—4 rounded, those of segments 5 and 
6 with sharp points. Telson with 3 pairs of lateral spines. Spines at apex of telson 
broken. 

Eye darkly pigmented, round. 

Stylocerite reaching end of first segment of antennular peduncle. First segment 
with 2 lateral spines. Second segment less than 0.5 length of first, with 1 spine. 
Third segment and flagella broken. Entire peduncle ‘‘about 0.5x length of anten- 
nal scale’? (Walker 1898). 

Basis of second antenna with sharp ventral spine, large blunt knob above. 
Scaphocerite broken, but broad. Flagella missing. 

Mouthparts dehydrated and fused in holotype. First and second maxillipeds 
with exopods. Third maxilliped without exopod, but with epipod, “‘reaching be- 
yond end of antennal scales’’ (Walker 1898). 

First pereopods strongly chelate, overreaching scaphocerite, with epipods. Fin- 
gers shorter than palm, with tuft of setae at tip. Propodus about 2x length of 
carpus, merus about equal to propodus. Ischium about 0.5 length of merus. 

Second pereopods chelate, with epipods. Fingers of chela shorter than palm. 
Carpus with 7 segments, the third of these the longest. Merus and ischium about 
equal, both about 0.5x length of carpus. 

Third pereopods with long, curved simple dactyls. Dactyls with a few long 
setae on curved margin, about 0.33 length of propodus. Carpus about 0.5 x 
propodus, with overhanging knob at articulation with propodus. Merus longer 
than propodus, with 2 spines. Ischium about 0.5 merus. Fourth and fifth per- 
eopods similar to third, all without epipods. Fourth merus with 4 lateral spines, 
fifth merus without spines. 

Pleopods dehydrated in holotype. Eggs small, numerous. Uropods broad, ovate. 

Type-locality.—**Puget Sound’’ (Walker 1898). 

Remarks.—The unique holotype of Heptacarpus herdmani has been broken 
and dehydrated so that many fine details cannot be observed. However, the lack 
of an exopod on the third maxilliped clearly shows that the species does not 
belong to the genus Euvalus. Indeed, Walker noted the lack of an exopod in his 
original description. 

Heptacarpus herdmani is related to H. paludicola Holmes, and H. flexus (Rath- 
bun) in having epipods on the first and second pereopods. Heptacarpus paludicola 
differs from H. herdmani in having 6-8 dorsal rostral spines and 2—4 ventral 
rostral spines, sharp points on the pleura of the fourth abdominal segment, and 
bifid dactyls with 5 spines on the third to fifth pereopods. Heptacarpus flexus has 
a very Slender rostrum with 4—5 dorsal spines and 5-8 ventral spines, a pro- 
nounced hump on the dorsal surface of the third abdominal segment, and a sub- 
orbital spine consisting of a prominent knob. 

One may ask who was responsible for the confusion between H. herdmani and 
the species of Eualus. Rathbun (1904:60) grouped Spirontocaris herdmani with 
species having both an exopod and an epipod on the third maxilliped and epipods 
on the first, second, and third pereopods. Specimens from the U.S. National 
Museum labelled §. herdmani in Rathbun’s handwriting belong to the new species 
of Eualus, not Heptacarpus (nor Spirontocaris, as now interpreted). Apparently, 
Rathbun missed Walker’s mention of the lack of exopods on the third maxillipeds, 
or assumed that he made an error in his description. 


‘VOLUME 96, NUMBER 1 3 


Fig. 1. Eualus lineatus: Dorsal and lateral views of female, carapace length 3.1 mm. 


AS now interpreted, the only known specimen of H. herdmani is the holotype 
from Puget Sound, either from Washington, U.S.A. or the Canadian side. No 
depth or substrate were mentioned for the specimen, but the long, simple dactyls 
suggest that the species lived on a soft bottom. Perhaps future dredging or sam- 
pling with SCUBA gear will discover more specimens of this poorly known shrimp. 


Eualus lineatus, new species 
Figs. 1-2 


Spirontocaris herdmani.—Rathbun, 1904:100—101 (in part). 
Eualus herdmani.—Holthuis, 1947:11 (in part)—Hobson and Chess, 1974:579.— 
Word and Charwat, 1976:103—104 (in part).—Butler, 1980: 197-198 (in part). 


Description.—Rostrum short, reaching second segment of antennular peduncle, 
Straight, with 3—6 dorsal teeth and 1-3 ventral teeth, all of dorsal margin toothed. 

Carapace smooth, dorsum straight. Small suborbital spine, moderate antennal 
spine with supporting carina, moderate pterygostomial spine. 

Abdomen smooth. Pleura of segments 1-3 rounded, pleura of fourth segment 
with weak posterolateral spine, fifth with strong posterolateral spine. Median 
dorsal margin of third segment strongly produced posteriorly. Telson with 3 pairs 
dorsolateral spines. 


4 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


0.5mm 


Fig. 2. Eualus lineatus: A, Anterior lateral view of female, carapace length 3.1 mm; B, Left third 
pereopod of female; C, Dacty] of left third pereopod of same female; D, Appendix masculina of male, 
carapace length 4.1 mm. 


Eyes moderately large, darkly pigmented. 

Stylocerite reaching end of first segment of antennular peduncle. First segment 
with 3 moderate dorsal spines. Second segment about 0.5 length of first, with 
2 strong spines. Third segment shorter than second, with 2 spines. Inner flagellum 
more than 2x length of outer. 

Basis of second antenna with sharp ventral spine, large blunt upper lobe. 
Scaphocerite oblong, shorter than carapace, lamella slightly longer than spine. 
Flagellum slightly exceeding body length. 

Mandible with slender incisor process, ending in small teeth. Molar process 
with spinules. Two-jointed palp present. First maxilla with lower endite slender, 
upper broad; palp faintly bilobed. Second maxilla with lower endite reduced, 
upper endite larger, bilobed; palp and scaphognathite well developed. First max- 
illiped with exopod and 2-jointed palp; epipod bilobed, caridean lobe very small; 
endites of coxa and basis separated by notch. Second maxilliped with exopod, 
podobranch, and epipod. Third maxilliped with exopod and epipod, exceeding 
antennular peduncle, stout and setose. 

First pereopod shorter than third maxilliped, stout, with epipod. Fingers of 
chela less than 0.5x length of palm. Carpus slightly shorter than palm. Merus 
about 2x length of carpus. Ischium less than 0.5x merus. Second pereopod 
chelate, longer than first pereopod, with epipod. Fingers shorter than length of 
palm. Carpus with 7 segments, the third and fourth the longest. Merus about 0.6 
length of carpus, slightly longer than ischium. Third pereopod slightly longer than 
second, with epipod. Dactyl stout, about 0.2 length of propodus, with 5 spines. 
Propodus with 16-19 spinules in 2—3 rows. Carpus about 0.4 length of propodus. 
Merus about same length as propodus, with 3 spines. Ischium less than 0.5x 
length of merus. Fourth pereopod about as long as third. Dactyl stout, similar to 
that of third pereopod. Propodus with 15 spinules in single row. Merus with 2-3 
spines. Fifth pereopod as long as fourth. Dactyl about 0.2 length of propodus, 


VOLUME 96, NUMBER 1 5 


with 4-5 small spines. Propodus with 14—22 spinules in | or 3 rows. Merus with 
0-1 spine. No epipods on pereopods 3-5. 

Second pleopod with appendix interna. Appendix masculina little more than 
half length of appendix interna, stout, apex truncated, with 8 long spinules. Uro- 
pods reaching end of telson, with long tooth on outer margin. 

Type-material—HOLOTYPE: female, total length 21.0 mm. 1.5 mi. SW of 
Gull Island, off Santa Cruz Island, California (33°56'00"N, 119°50’55”W), 89 m, 
mud and sponge bottom, 8 Nov 1941, Velero IIT sta. 1435-41, AHF type number 
4129.—Paratype: female. San Juan Islands, Washington, summer 1930, John C. 
Queen, collector, USNM.—Paratypes: female, carapace length 3.1 mm, west 
shore Jesse Island, British Columbia (49°12'’N, 123°57'W), 30 Apr 1972; female, 
carapace length 3.4 mm, Boca del Infierno Bay, Nootka Sound, British Columbia 
(49°38’N, 126°37’W), 21 May 1973, National Museum of Canada. 

Other material.—41 specimens from 24 stations, AHF and USNM, Naha Bay, 
Alaska; coast of British Columbia, Puget Sound, off Depoe Bay, Oregon; south- 
ern California from Redondo Beach and off Long Beach; Santa Cruz, Santa Rosa, 
and Santa Catalina Islands; Bahia San Gabriel, Isla Espiritu Santo, Gulf of Cal- 
ifornia, Mexico; 17-140 m, among rocks, kelp, sand or mud; on wreck (MKW). 
34 specimens from Burnaby Narrows, Queen Charlotte Islands (52°22'N, 131°21’W) 
to Jesse Island (49°12’N, 123°57'W), intertidally on sand and gravel to 232 m on 
mud, 1928-1973 (THB). 

Records in literature.—Sitka, Alaska to Puget Sound (Butler 1980); Santa Mon- 
ica Bay, Palos Verdes Peninsula, Santa Catalina Island, and Point Loma, Cali- 
fornia (Word and Charwat 1976). 

Color in life. —Red diagonal lines on carapace and first and second abdominal 
segments, red spots and blotches on third to sixth abdominal segments, telson, 
uropods, and protopodites of pleopods, smaller red spots on all anterior append- 
ages, including eye stalk and pereopods (Butler 1980, color plate 1C). 

Parasites.—Parasitized by Bopyroides hippolytes (Butler, 1980). An unidenti- 
fied rhizocephalan on a specimen from Neah Bay, Washington. 

Remarks.—Eualus lineatus is related to E. avinus (Rathbun), E. pusiolus (Kr@yer), 
and E. berkeleyorum Butler in having epipods on the first 3 pereopods. In Eualus 
avinus the rostrum is arched over the eye, and bears 12—14 upper teeth and 1-3 
ventral teeth. The dactyl of the third pereopod is slender and simple. Eualus 
pusiolus has a straight rostrum shorter than the second segment of the antennular 
peduncle, with 2-5 upper teeth and at most | lower tooth. The dactyl of the third 
pereopod is stout. Eualus berkeleyorum has a straight rostrum with 8-11 dorsal 
teeth and 2—5 ventral teeth. The dactyls of its third pereopods are slender and 
simple. It does not have a ventral spine on the pleura of the fourth abdominal 
segment. 

Most specimens of E. lineatus have been taken by trawls. One specimen was 
collected by a SCUBA diver on the wreck of the Olympic, off Long Beach, 
California. A first stage larva of the species has been hatched in the laboratory 
(Butler 1980). The species is eaten by rockfishes (Sebastes spp.) off Santa Cat- 
alina Island, California (Hobson and Chess 1974). 

Etymology.—The name of the species refers to the red lines on the carapace 
and body of the shrimp. 


6 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Literature Cited 


Butler, T. H. 1980. Shrimps of the Pacific coast of Canada.—Canadian Bulletin of Fisheries and 
Aquatic Sciences 202, 280 pp. 

Hobson, E. S., and J. R. Chess. 1976. Trophic interactions among fishes and zooplankters near 
shore at Santa Catalina Island, California.—United States National Marine Fisheries Service 
Fishery Bulletin 74(3):567—598. 

Holmes, S. J. 1900. Synopsis of California stalk-eyed Crustacea.—Occasional Papers of the Cali- 
fornia Academy of Sciences 7:1—262, pls. 1-4. 

Holthuis, L. B. 1947. The Hippolytidae and Rhynchocinetidae collected by the Siboga and Snellius 
expeditions with remarks on other species. The Decapoda of the Siboga Expedition Part IX.— 
Siboga Expeditie Monographie 39a: 1-100. 

Rathbun, M. J. 1904. Decapod crustaceans of the northwest coast of North America.—Harriman 
Alaska Expedition, Series 10, 210 pp. 

Walker, A. O. 1898. Crustacea collected by W. A. Herdman in Puget Sound, Pacific coast of North 
America, September, 1897.—Proceedings and Transactions of the Liverpool Biological Society 
12:268—287. 

Wicksten, M. K. 1979. Zoogeographical affinities of the broken back shrimp (Caridea: Hippolytidae) 
of western South America.—Proceedings of the International Symposium on Marine Biogeog- 
raphy and Evolution in the Southern Hemisphere. Vol. 2. New Zealand Division of Scientific 
and Industrial Research Information Series 137:627-634. 

Word, J. Q., and D. Charwat. 1976. Invertebrates of southern California coastal waters. II. Natan- 
tia.—El Segundo: Southern California Coastal Water Research Project. 238 pp. 


(MKW) Department of Biology, Texas A&M University, College Station, Tex- 
as 77843; (THB) Pacific Biological Station, Nanaimo, British Columbia V9R 5K6, 
Canada. 


PROC. BIOL. SOC. WASH. 
96(1), 1983, pp. 7-28 


HYDROIDA FROM ESTUARIES OF SOUTH CAROLINA, 
USA: FAMILIES SERTULARIIDAE AND 
PLUMULARIIDAE 


Dale R. Calder 


Abstract.—Six species of Sertulariidae and five species of Plumulariidae were 
identified in collections of hydroids from estuaries of South Carolina. Sertularia 
exigua Allman, 1877, is synonymized with Dynamena cornicina McCrady, 1859, 
and Aglaophenia rigida Allman, 1877, is placed in synonymy with A. trifida L. 
Agassiz, 1862. Aglaophenia trifida occurs in two distinct colony forms in estuaries 
of South Carolina, but the two variants are not regarded as separate taxa. Obelia 
marginata Allman, 1877, from the Atlantic Ocean and Campanularia marginata 
Bale, 1884, from Australia belong to the genus Thyroscyphus Allman, 1877. Thy- 
roscyphus balei, new name, is proposed as a replacement name for the junior 
secondary homonym 7. marginatus (Bale, 1884). 

The assemblage has a distinct warm water affinity. All 11 species range at least 
as far south as the Caribbean, while but four occur north of the Carolina coast 
on substrates other than pelagic Sargassum. Only two of the species, Sertularella 
conica and Aglaophenia rigida, are restricted to the western Atlantic. No rep- 
resentatives of either family were collected below salinities of 15% in the study 
area. 


Studies by McCrady (1859) and Fraser (1912) form the basis for much of the 
information presently available on hydroids of estuarine and nearshore areas 
between Cape Hatteras, North Carolina, and Cape Canaveral, Florida. Other 
records of hydroids from the Carolinian region are scattered in various taxonomic 
papers covering broad geographic areas, such as those of Allman (1877), Jader- 
holm (1896), Nutting (1900, 1904, 1915), Stechow (1923), Fraser (1943, 1944) and 
Vervoort (1972). Additional data are given in investigations of fouling commu- 
nities (McDougall 1943; Dean and Bellis 1975; Sutherland 1977, 1981; Sutherland 
and Karlson 1977; Karlson 1978), and faunal studies (Pearse 1936; Pearse and 
Williams 1951; Stephenson and Stephenson 1952; Wells 1961; Wells, Wells, and 
Gray 1964; Cain 1972; Howard and Frey 1975), as well as miscellaneous other 
publications (Verrill 1872; Fewkes 1881; Brooks 1886; Mayer 1910a, b; Sandifer 
et al. 1974; Calder 1976; Calder and Hester 1978). Despite its dated systematics, 
Fraser’s (1944) monograph continues to be the most comprehensive source of 
information on hydroids of the region. 

Hydroids occur from marine to freshwater environments and are an important 
constituent of epibenthic assemblages in estuaries. This paper is a systematic 
account of the species belonging to the families Sertulariidae and Plumulariidae 
found in estuaries of South Carolina. 


Materials and Methods 


Hydroids were sorted from benthic samples collected between 1973 and 1978 
at 130 stations in estuaries of South Carolina. These stations encompassed the 


8 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Table 1.—Locations, depths, substrate types, and observed bottom salinity ranges for stations 
where hydroids of the families Sertulariidae and Plumulariidae were collected in estuaries of South 
Carolina. 


Depth Salinity 
Station Location N WwW (m) Substrate (%o) 
LRA7 Little River Inlet 33°51'40" 78°32'31" 4 sand, shell 33 
LRA8 Little River Inlet 33°52'00" 78°32’ 48" y sand, shell 33 
MI07 Murrells Inlet 33°30'00" 79°01'34" 3 sand, shell 34 
MI09 Murrells Inlet 33°33'12” 79°01'43” 2 sand, shell 34 
M110 Murrells Inlet B363 050 79°01'44" 3 sand, shell 34 
MI12 Murrells Inlet 33°33'16" 79°01'08" 1 sand, shell 34 
MI13 Murrells Inlet 33°33'28" 79°01'35" y, sand, shell 34 
MI15 Murrells Inlet 33°33'37" 79°01'24" 1 sand, shell 34 
MI16 Murrells Inlet 33°33’ 18” 79°02'02" 3 sand, shell 34 
MI17 Murrells Inlet 33°33'08" 79°02’ 14” 1 silty sand, shell 34 
YBO02 Winyah Bay 33°13'21" 79°11'14" 5 sand, shell — 
NS00 North Santee River 33°09'00" 79° 14'09” 0 sand, shell = 
NSO] North Santee River 33°08'34" 79°14'48" 3 sand, shell = 
NB02 North Santee River 33°10'00" 79°14'30" 1 sand, shell —_— 
BBOl1 Bulls Bay SPS 2 79°36'57" 4 mud, shell 34 
BBO2 Sewee Bay BS a” 79°38'15" D; mud, shell 34 
SB03 Sewee Bay 32°56'51” 79°38'33" 2 mud, shell 32-35 
SB04 Sewee Bay 32°56'47" 79°38'30" 3 mud, shell 32-35 
SBOS5 Sewee Bay 32°56'35” 79°39'07" 4 mud, shell 32-35 
SB07 Sewee Bay 32°56'24" 79°39'00" 2 mud, sand, shell 31-35 
SB08 Sewee Bay 32°56'17" 79°38' 45” 2 mud, shell 31-35 
SB10 Sewee Bay 32°55'49" 79°39'25" 4 mud, shell 31-34 
SB13 Sewee Bay Se) D1 79°38'29" 4 mud, shell 31-35 
SB15 Sewee Bay 32°55'23" 79°39'37" 4 mud, shell 31-34 
SB18 Sewee Bay BPS) A! 79°38'27" y mud, shell 30-35 
B003 Bulls Bay 32°55.9’ 79°36.2' 5 mud, sand 28-34 
B002 Prices Creek 32°54.2’ 79°40.7' 8 sand, shell 29-34 
CI01 Capers Inlet 32°51'30" 79°42'35" 3 sand, shell 34 
B00] Inlet Creek 32°47.5’ 79°49.5' 4 sand, shell 23-33 
CH00 Charleston Harbor 32°45'20" 79°51'40" 20 sand, shell — 
J003 Charleston Harbor 32°44.9' 79°51.6' 10 mud, shell 22-33 
J002 Charleston Harbor 32°47.1' TES 3.2 3 mud 16—28 
JO01 Charleston Harbor 32°45.4’ 79°55.1 7 mud, sand 16-25 
CH02 Charleston Harbor 32°46’ 18” 79°53'17" 12 mud, shell — 
KPO1 Stono Inlet 32°37'25" 79°59'26" 10 sand, shell 29 
KPO02 Stono River 32°38'03” 80°00'32” 10 sand, shell 28 
KP04 Stono River 32°40'29" 80°00’ 10” 7 sand, shell 24 
KP07 Kiawah River 32°37 '29" 80°04'33” 6 sand, shell 25 
KP08 Kiawah River 32°36'54” 80°06’ 43” 2 mud, shell 24 
E001 Wadmalaw Sound 32°41.2' 80°10.4’ 7 sand, shell, mud 17-27 
DE10 North Edisto River 32°39'06" 80°15'25” 11 sand, shell 22-33 
E003 North Edisto River 32°38.8' 80°15.7' 7 sand, shell 19-3] 
E002 Toogoodoo Creek 32°41.3’ 80°17.3’ 4 sand, shell, mud 21-26 
DE04 North Edisto River 32°37'41” 80°16'34” 7 sand, mud, shell 22-33 
DEO05 North Edisto River BD Bila 80°16’ 16" 10 pebbles 21-34 
DE06 North Edisto River 32°36'53” 80°15'55” 10 sand, shell 27-31 
E005 Steamboat Creek 32°36.2’ 80°17.7' 7 sand, mud, shell 23-29 
DE07 North Edisto River 32°36'33” 80°15’10” 9 sand, shell 27-31 
DE08 North Edisto River 32°36'21” 80°14'00" 8 sand 24-34 
E006 North Edisto River 32°36.5’ 80°14.8' 8 sand 25-30 
E007 North Edisto River 32°35.9! 80°13.5' 8 mud 26-3 1 


VOLUME 96, NUMBER | 9 


Table 1.—Continued. 


Depth Salinity 
Station Location N WwW (m) Substrate (%o) 
ACO1 Adams Creek 32°35'48" 80°12’22” 3 sand, shell — 
E008 North Edisto River 32°33.6' 80°10.7' 10 sand, shell 29-31 
DEO! Dawho River 32°37'25” 80°17'05” 4 mud 19-29 
D004 South Edisto River 32°29.7' 80°21.2' 7 sand 24-33 
H003 St. Helena Sound 32°30.9' 80°27.9’ 5 sand, mud, shell 21 
BRO8 Beaufort River 32°24'46" 80°40'53” 6 sand, shell = 
P002 Colleton River 32°16.2’ 80°43.7’ 6 mud, sand 27-32 
P00! Port Royal Sound 32°16.2’ 80°48.5’ 8 sand, mud, shell 24-31 
P006 Chechessee River 32°17'30" 80°45’00" 9 sand 30 
G001 Calibogue Sound 32°10.9' 80°47.8’ 7 sand, mud, shell 25-30 


region from Little River Inlet in the northeast to the Wright River in the south- 
west, and extended from the mouth to the head of various estuarine systems in 
the state. Representatives of the families Sertulariidae and Plumulariidae were 
found at 61 of these stations (Table 1). Most specimens were obtained in epifaunal 
collections made with modified oyster dredges and otter trawls, but a few were 
sorted from infaunal samples taken using Petersen grabs. Collections were pre- 
served in 10% neutralized formaldehyde. 

Bottom water samples, taken in conjunction with most benthic collections, 
were obtained with Van Dorn bottles and analyzed for salinity in the laboratory 
with a Beckman RS7B induction salinometer. 

Hydroids occurring on pelagic Sargassum are allochthonous to South Carolina 
estuaries and were not included in this study. The synonymy list for each species 
includes published records from estuaries between Cape Hatteras, North Caro- 
lina, and Cape Canaveral, Florida. Descriptions and illustrations are based en- 
tirely on specimens from estuaries of South Carolina. The terminology employed 
largely follows that used in the monograph on South African hydroids by Millard 
(1975). A reference collection has been deposited in the Department of Inverte- 
brate Zoology, Royal Ontario Museum. 


Systematic Account 


Family Sertulariidae Fleming, 1828 
Dynamena Lamouroux, 1812 
Dynamena cornicina McCrady, 1859 
Figs. 1, 2 


Dynamena cornicina McCrady, 1859:204.—Calder, 1976:169.—Calder and Hes- 
ter, 1978:91. 

Sertularia cornicina.—Verrill, 1874:408, 411, 733.—Fraser, 1912:374, fig. 38; 1944: 
279, fig. 266. 

Sertularia exigua Allman, 1877:24, pl. 16, figs. 7, 8. 

Sertularia complexa Clarke, 1879:245, pl. 4, figs. 26-28. 

Sertularia moluccana Pictet, 1893:50, pl. 2, figs. 42, 43. 

Desmoscyphus palkensis Thornely, 1904:119, pl. 2, figs. 7A, B. 

Sertularia densa Stechow, 1919:93, fig. J1. 

Sertularia cornicina f. pinnata Jarvis, 1922:339. 


10 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Collection records.—Sta. MI07, dredge, 29 May 1975. Sta. MI09, dredge, 29 
May 1975. Sta. MI10, dredge, 29 May 1975. Sta. MI12, dredge, 29 May 1975. 
Sta. MI13, dredge, 29 May 1975. Sta. MI15, dredge, 29 May 1975. Sta. MI16, 
dredge, 29 May 1975. Sta. BBO2, dredge, 11 Aug 1977. Sta. SB03, dredge, 22 
Aug 1977. Sta. SB04, dredge, 25 July 1978. Sta. SBO7, dredge, 22 Aug 1977. Sta. 
SB10, dredge, 22 Aug 1977, 25 July 1978. Sta. SB13, dredge, 25 July 1978. Sta. 
SB15, dredge, 22 Aug 1977. Sta. BO03, dredge, 12 Aug 1974, 22 Oct 1974, 13 Oct 
1975. Sta. BOO2, dredge, 23 Oct 1973, 15 Apr 1974, 12 Aug 1974, 22 Oct 1974, 13 
Oct 1975, 5 Jan 1976. Sta. BOOI, dredge, 16 Oct 1974, 2 Apr 1976. Sta. JO03, 
dredge, | Aug 1975, 1 Apr 1976. Sta. KP07, dredge, 15 Mar 1977. Sta. KP08, 
dredge, 16 Mar 1977. Sta. DE04, dredge, 21 Nov 1977, 17 Oct 1978. Sta. DEOS, 
dredge, 17 Oct 1978. Sta. DE07, dredge, 21 Nov 1977. Sta. E008, dredge, 2 Oct 
1974. Sta. D004, dredge, 9 Jan 1974. Sta. BRO8, dredge, 27 Mar 1974, 8 Oct 1975. 
Sta. P002, dredge, 4 Oct 1973, 5 Apr 1974, 7 Aug 1974, 8 Oct 1974. Sta. POO, 
dredge, 4 Oct 1973, 7 Aug 1974, 8 Oct 1974. Sta. P006, trawl, 7 Aug 1974. Sta. 
G001, dredge, 4 Oct 1973, 7 Aug 1974, 8 Oct 1974, 8 Apr 1975, 8 Oct 1975. 

Description.—Unbranched colonies reaching 2 cm high; color in life bright 
yellow. Hydrocaulus monosiphonic, straight, arising from a creeping hydrorhiza; 
basal part athecate, length variable but usually about | mm long, articulating with 
upper thecate part by an oblique hinge-joint; thecate part divided by oblique 
nodes, internodes 491-784 ym long, each with an opposite pair of frontally-placed 
hydrothecae; thecate part of a few hydrocauli having an irregularly-placed athe- 
cate internode marked by an oblique hinge-joint distally. Hydrothecae cylindrical, 
distal half curved outward, contiguous frontally, not touching across back of 
internode; abcauline wall concave except for an occasional basal bulge, 293-386 
ym long; contiguous part of adcauline wall straight, 246-363 um long, free part 
convex, 199-293 wm long; diameter at base of hydrotheca 129-188 wm. Hy- 
drothecal orifice oval, 129-158 um wide from adcauline to abcauline wall; margin 
with 2 prominent lateral teeth and a smaller median adcauline tooth, occasionally 
renovated. Operculum consisting of a large abcauline and a smaller adcauline 
valve divided by a median line. Perisarc of abcauline wall expanded just below 
margin, but intrathecal teeth lacking. 

Gonothecae oval, resembling chinese lanterns with about 6 rounded, transverse 
ridges, 680 um wide, 1000 um long from base to orifice, arising by short pedicels 
from hydrorhiza. Walls round in cross-section, orifice round, 365 «wm in diameter, 
at end of short collar. 

Remarks.—Charleston Harbor, South Carolina, is the type-locality of this cir- 
cumglobal hydroid. According to Nutting (1904), McCrady’s (1859) types of this 
species were destroyed by fire in Charleston during the American Civil War. I 
was unable to locate any of McCrady’s hydrozoan material at the Charleston 
Museum. 

Sertularia exigua, originally described by Allman (1877) from Cape Fear, North 
Carolina, is regarded as a synonym of D. cornicina. Allman’s types of S. exigua 
could not be located at the Museum of Comparative Zoology where they were 
originally deposited (Nutting 1904), but his descriptions and illustrations of the 
species are indistinguishable from D. cornicina. The back of the hydrothecal 
pairs, rather than the front, is shown in Allman’s drawings of the species. 

Dynamena cornicina is often a substrate for the hydroid Hebella scandens 


~ VOLUME 96, NUMBER 1 11 


(Bale 1888), and it was probably hydrothecae of the latter that McCrady (1859) 
presumed, with considerable reservation, to be gonothecae. 

Known range.—Western Atlantic: Massachusetts to Brazil; Bermuda. Else- 
where: Circumglobal, tropical and temperate waters. 


Dynamena quadridentata (Ellis and Solander, 1786) 
Fig. 3 


Sertularia quadridentata Ellis and Solander, 1786:57, pl. 5, fig. G. 

Pasythea (Sertularia) quadridentata.—Lamouroux, 1812:183. 

Pasythea quadridentata.—Lamouroux, 1816:156.—Fraser, 1912:372, fig. 36. 
Pasythea nodosa Hargitt, 1908:114, figs. 13-15. 

Pasya quadridentata.—Stechow, 1923:166. 

Pasya elongata Stechow and Muller, 1923:469, pl. 27, fig. 8. 

Dynamena gibbosa Billard, 1924:650, fig. 2G. 

Dynamena quadridentata.—Billard, 1925:194, fig. 42. 

Dynamena quadridentata var. elongata Billard, 1925:195, fig. 43A, B, C, D. 
Dynamena quadridentata var. nodosa Billard, 1925:197, fig. 43E. 


Collection record.—Sta. J003, dredge, | Apr 1976. 

Description.—Hydrorhiza with distinct internal ridges of perisarc. Colonies 4.5 
mm high; hydrocaulus monosiphonic, straight, unbranched; basal portion athe- 
cate, short, with an oblique hinge-joint distally; thecate part divided either by 
oblique and frequently indistinct nodes, or by oblique hinge-joints. Internodes 
variable in length with 1-3 opposite pairs of frontally-situated hydrothecae; hy- 
drothecal pairs usually contiguous for a varying distance in front, not touching 
across back of internode. Base of hydrotheca above abutting adcauline wall of 
hydrotheca below when 2 or more pairs of hydrothecae present per internode. 
Hydrothecae varying from nearly cylindrical to somewhat tumid, narrowest just 
below margin; abcauline wall 187-281 um long; solitary pair, or proximal pairs 
of a group curved outward with axes oblique to internode, adcauline wall convex, 
abcauline wall concave except for basal bulge; distal pair of a group with adnate 
portions of adcauline wall straight, axes parallel to internode, margin curved 
abruptly outward, abcauline wall concave. Hydrothecal margin oval with 2 prom- 
inent lateral teeth and a smaller median adcauline tooth. Operculum consisting 
of a large abcauline and a smaller adcauline valve. Intrathecal teeth present. 

Gonothecae absent. 

Remarks.—This species is common on pelagic Sargassum, but was collected 
from benthic algae at only one station in the study area. 

Known range.—Western Atlantic: North Carolina to Brazil; Bermuda; north- 
ward to Massachusetts on floating Sargassum. Elsewhere: Circumglobal, tropical 
and warm temperate waters. 


Sertularella Gray, 1848 
Sertularella conica Allman, 1877 
Fig. 4 
Sertularella conica Allman, 1877:21, pl. 15, figs. 6, 7.—Fraser, 1912:373, fig. 37. 
Sertularella gayi.—Verrill, 1900:571; 1907:319, fig. 170 [not Sertularella gayi (La- 
mouroux, 1821)]. 


12 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Sertularella inconstans.—Calder and Hester, 1978:91 (not Sertularella inconstans 
Billard, 1919). 


Collection record.—Sta. CH00, trawl, 22 May 1974. 

Description.—Unbranched, sympodial colonies reaching | cm high. Hydrocau- 
lus monosiphonic, geniculate, divided into internodes by oblique nodes sloping 
alternately in opposite directions, an annulation often evident above each node; 
perisarc thick. Internodes variable in length, first very short; each internode with 
a distal hydrotheca; diameter at nodes 135-181 um. Hydrothecae alternate, slen- 
der, flask-shaped, widest at juncture of adcauline wall and internode, narrowest 
just below margin, walls with 4-8 distinct annulations; adcauline wall convex, 
length adnate 211-257 um, length free 293-445 um; abcauline wall slightly convex 
basally, slightly concave distally, 398-562 um long. Hydrothecal margin with 4 
distinct, equal teeth, operculum of 4 valves; aperture quadrate or irregularly oval, 
187-211 wm wide from adcauline to abcauline tooth; 3 well-developed internal 
teeth, 2 additional smaller ones occasionally evident. 

Gonothecae absent. 

Remarks.—This species is very similar to Sertularella robusta Coughtrey, 1876, 
as described by Pennycuik (1959), Ralph (1961), Blanco (1968), Vervoort (1972), 
Watson (1973), and Vervoort and Vasseur (1977). The latter species occurs over 
a wide area in the southern hemisphere. Sertularella conica has been observed 
in Bermuda (Calder, unpublished) and was recorded earlier from Bermuda by 
Verrill (1900, 1907) as Sertularella gayi (Lamouroux, 1821). Bennitt’s (1922) rec- 
ord of S. tenella (Alder, 1856) from Bermuda may also have been based on this 
species. Sertularella conica is readily distinguished from S$. gayi in having internal 
teeth below the margin of the hydrotheca. Fraser’s (1913, 1944) accounts of this 
species from the Canso Banks, Nova Scotia, as well as those from the eastern 
Pacific (Fraser 1937, 1948), do not correspond with S. conica. Specimens iden- 
tified as S. conica from the Tortugas by Gemerden-Hoogeveen (1965) also appear 
to be based on material of another species. 

Sertularella conica was collected once in estuaries of South Carolina. Speci- 
mens were found growing on hydrocauli of Ag/aophenia trifida taken at the en- 
trance of Charleston Harbor. 

Known range.—Western Atlantic: North Carolina to the Caribbean Sea; Ber- 
muda. 


Sertularia Linnaeus, 1758 
Sertularia distans Lamouroux, 1816 
Fig. 5 


Sertularia distans Lamouroux, 1816:191. Not Dynamena distans Lamouroux, 
1816:180, pl. 5, figs. la, lb [=Dynamena pumila (Linnaeus, 1758)]. 

Sertularia gracilis Hassall, 1848:2223. 

Dynamena gracilis. —Marktanner-Turneretscher, 1890:240 (part). 

Sertularia pourtalesi Nutting, 1904:59, pl. 5, fig. 5. 

Sertularia stookeyi Nutting, 1904:59, pl. 5, figs. 6, 7.—Fraser, 1912:375, fig. 39.— 
Calder, 1976:169.—Calder and Hester, 1978:91. 

Sertularia heterodonta Ritchie, 1909a:79, fig. 4 [not Dynaimena heterodonta (Sar- 
vis, 1922)]. 


VOLUME 96, NUMBER 1 13 


Sertularia distans var. garcilis Billard, 1912:465. 
Tridentata heterodonta.—Stechow, 1923:205. 
Tridentata pourtalesi.—Stechow, 1923:205. 
Tridentata stookeyi.—Stechow, 1923:205. 
Tridentata gracilis.—Stechow, 1923:208, fig. G. 
Sertularia distans gracilis Millard, 1964:49. 
Tridentata distans.—Hirohito, 1969:23, fig. 16. 


Collection records.—Sta. BBO1, dredge, 11 Aug 1977. Sta. BBO2, dredge, 11 
Aug 1977. Sta. SB03, dredge, 22 Aug 1977, 28 Apr 1978, 25 July 1978. Sta. SB04, 
dredge, 22 Aug 1977. Sta. SB05, dredge, 22 Aug 1977. Sta. SBO7, dredge, 22 Aug 
1977. Sta. SB10, dredge, 22 Aug 1977. Sta. SB15, dredge, 22 Aug 1977, 25 May 
1978. Sta. BOO3, dredge, 22 Oct 1974. Sta. B0O2, trawl, 23 Apr 1973, 23 July 1973, 
23 Oct 1973; dredge, 14 Jan 1974, 15 Apr 1974, 12 Aug 1974, 22 Oct 1974, 16 Apr 
1975, 13 Oct 1975, 5 Jan 1976. Sta. BOO], dredge, 9 Oct 1973, 17 Jan 1974, 12 
Apr 1974, 15 Aug 1974, 16 Oct 1974, 13 Oct 1975, 2 Jan 1976, 2 Apr 1976. Sta. 
J003, Petersen grab, | Oct 1973; dredge, 1 Oct 1973. Sta. CH02, dredge, 10 Oct 
1973. Sta. J0OO2, dredge, 14 Oct 1975. Sta. KPO1, dredge, 14 Mar 1977. Sta. KP02, 
dredge, 14 Mar 1977. Sta. KP04, dredge, 14 Mar 1977. Sta. KPO7, dredge, 15 
Mar 1977. Sta. DE10, dredge, 7 Mar 1978. Sta. E003, dredge, 2 Apr 1974, 2 Aug 
1974. Sta. E002, trawl, 15 Sep 1973. Sta. DE04, dredge, 15 Dec 1977. Sta. DEOS, 
dredge, 21 Nov 1977, 17 Oct 1978. Sta. DE06, dredge, 21 Nov 1977, 13 June 
1978. Sta. E005, dredge, 7 Jan 1974. Sta. DEO7, dredge, 21 Nov 1977, 15 Dec 
1977. Sta. DE08, dredge, 7 Mar 1978, 17 Oct 1978. Sta. E006, trawl, 7 Aug 1973; 
dredge, | Feb 1974. Sta. E007, dredge, 5 Aug 1974, 9 Oct 1975, 11 Jan 1976. Sta. 
E008, trawl, 9 Apr 1973, 2 July 1973, 5 Sep 1973; dredge, 2 Oct 1973, 7 Jan 1974, 
3 Apr 1974, 2 Oct 1974. Sta. D004, dredge, 3 Oct 1973, 9 Jan 1974. Sta. BRO8, 
dredge, 8 Oct 1975. Sta. P002, dredge, 5 Apr 1974, 8 Oct 1974. Sta. POOI1, dredge, 
5 Apr 1974. Sta. GOOI, dredge, 8 Oct 1974. 

Description.—Colonies up to 2 cm high; usually unbranched but occasionally 
with one or more branches; branch arising either from an apophysis inserting just 
below a hydrotheca or from renovation of an old hydrotheca. Hydrocaulus mon- 
osiphonic, straight, arising from a creeping hydrorhiza; basal part of variable 
length but usually less than 500 um long, consisting of 1 or 2 athecate internodes 
marked by oblique hinge-joints; thecate part divided by slightly oblique and often 
obscure nodes, internodes 398-620 um long, each with an opposite pair of fron- 
tally-placed hydrothecae; thecate part of some hydrocauli having one or more 
irregularly-placed athecate internodes with oblique hinge-joints distally. Hydro- 
thecae horn-shaped, curved outward; pair usually contiguous but occasionally 
separated frontally, not touching across back of internode; abcauline wall con- 
cave, 181-222 wm long; adcauline wall 246-281 um long, length contiguous 0- 
105 wm, diameter at base of hydrotheca 64-117 wm. Hydrothecal orifice oval, 
70—94 ym wide from adcauline to abcauline wall; margin with 2 prominent lateral 
teeth and a small median adcauline tooth. Operculum consisting of an abcauline 
valve and an adcauline valve with a median line. Intrathecal teeth lacking. Hy- 
dranth with an abcauline caecum. 

Gonothecae oval, with about 4 faint, rounded ridges, maximum diameter 351- 
433 wm, length from base to orifice 749-842 um, arising by short pedicels from 


14 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


6 


Figs. 1-8. Family Sertulariidae. 1, Dynamena cornicina, hydrothecae, station P001; 2, Dynamena 
cornicina, gonotheca, station P001; 3, Dynamena quadridentata, hydrothecae, station J003; 4, Ser- 
tularella conica, hydrothecae, station CH00; 5, Sertularia distans, hydrothecae and gonotheca, station 
E002; 6, Sertularia marginata, hydrothecae, station D004; 7, Sertularia marginata, gonotheca, station 
D004; 8, Thyroscyphus marginatus, hydrothecae, station B002, scale bars = 500 um. 


VOLUME 96, NUMBER | 15 


hydrocaulus at bases of hydrothecae. Walls oval or nearly round in cross-section; 
orifice round, 211-252 um in diameter, at end of a short collar; submarginal teeth 
present. 

Remarks.—Specimens from South Carolina were compared with Nutting’s (1904) 
type—material of S. stookeyi from the Great Bahama Banks (USNM 19710) and 
are inseparable from them morphologically. Sertularia stookeyi has been included 
in the synonymy of S. distans Lamouroux, 1816 by Cornelius (1979). 

Sertularia distans is widespread in estuaries of South Carolina in salinities 
above 21%c. It frequently occurs in dense masses on the axes of dead specimens 
of the octocoral Leptogorgia virgulata. 

Known range.—Western Atlantic: Massachusetts to Brazil; Bermuda. Else- 
where: Circumglobal, tropical and temperate waters. 


Sertularia marginata (Kirchenpauer, 1864) 
Figs. 6, 7 


Dynamena marginata Kirchenpauer, 1864:13, fig. 8. Not Sertularia marginata 
Allman, 1877:23, pl. 16, figs. 1, 2 (=Sertularia striata Totton, 1930). 

Sertularia flosculus Thompson, 1879:104, pl. 17, fig. 2. 

Sertularia amplectens Allman, 1885:141, pl. 16, figs. 3, 4. 

Desmoscyphus pectinatus Allman, 1888:71 (part). 

Desmoscyphus gracilis Allman, 1888:71, pl. 34, figs. 2, 2a—c. 

Desmoscyphus inflatus Versluys, 1899:42, figs. 11-13. 

Sertularia inflata.—Jaderholm, 1903:286. 

Sertularia versluysi Nutting, 1904:53, pl. 1, figs. 4~9.—Fraser, 1912:375, fig. 40. 

Sertularia pluma Hartlaub, 1905:661. 

Sertularia marginata.—Bale, 1913:125 (part).—Calder, 1976:169.—Calder and 
Hester, 1978:91. 

Amphisbetia marginata.—Stechow, 1921:258. 

Tridentata flosculus.—Stechow, 1923:204. 

Tridentata inflata.—Stechow, 1923:205. 

Sertularia marginata f. typica Vannucci, 1949:248. 

Sertularia marginata f. laxa Vannucci, 1949:248, pl. 3, fig. 46. 


Collection records.—Sta. J003, dredge, | Oct 1973. Sta. J002, dredge, 14 Oct 
1975. Sta. JOO1, trawl, 8 Oct 1973. Sta. D004, dredge, 3 Oct 1973. Sta. H003, 
dredge, 10 Oct 1975. 

Description.—Hydroids up to 6.5 cm high, arising from a tangled hydrorhiza. 
Hydrocaulus straight, monosiphonic, alternately branched, perisarc thick; basal 
part athecate, length variable but usually several mm long, occasionally with one 
or more transverse or slightly oblique nodes, separated from upper thecate part 
by an oblique hinge-joint; thecate part divided by rather indistinct transverse 
nodes, internodes short, each with an apophysis and an axillary hydrotheca ba- 
sally and a pair of subopposite hydrothecae distally, hydrothecal pair separated 
front and back. Apophyses given off alternately from opposite sides of hydro- 
caulus, separated from hydroclade by a transverse node, node sometimes ob- 
scure. Hydrocladia unbranched, reaching | cm long, directed upward; first inter- 
node short, athecate, node transverse proximally, an oblique hinge-joint distally; 
remaining internodes each with one pair of frontally-placed hydrothecae, nodes 


16 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


oblique, often indistinct or entirely obliterated. Hydrothecae in opposite pairs, 
pair occasionally separated but usually contiguous frontally, not touching across 
back of internode; each hydrotheca expanded basally, turned abruptly outward 
and slightly upward distally, tapering towards margin; abcauline wall concave, 
176-222 um long with a prominent intrathecal septum; contiguous part of adcau- 
line wall straight, length contiguous 0-211 um, free part convex, 129-304 um 
long. Hydrothecal orifice oval, margin with very small median adcauline tooth 
and 2 large, pointed lateral teeth appearing unequal in size due to angle of ap- 
erture. Operculum consisting of a large abcauline and a smaller adcauline valve 
divided by a median line. 

Gonothecae resembling Chinese lanterns, 760 ~m wide, 1400 um long from 
base to orifice, arising by short pedicels from hydrocaulus just above axillary 
hydrothecae. Walls oval in cross-section, with 5—6 transverse ridges; orifice hemi- 
spherical, bounded by 2 spines; spines horn-shaped, 255-350 um long. 

Remarks.—Van Gemerden-Hoogeveen (1965) recognized that Sertularia inflata 
(Versluys, 1899) was very difficult to separate from S. marginata, but she be- 
lieved that they could be distinguished on the basis of differences in gonotheca 
arrangement and shape as well as hydrotheca shape. However, I have followed 
Billard (1925), Vannucci Mendes (1946), Ralph (1961), Mammen (1965a), Millard 
and Bouillon (1974), Millard (1975), Garcia et al. (1980), and others in regarding 
S. inflata as a synonym of S. marginata. 

Known range.—Western Atlantic: North Carolina to Brazil; Bermuda; north- 
ward to Massachusetts on floating Sargassum. Elsewhere: Circumglobal, tropical 
and subtropical waters. 


Thyroscyphus Allman, 1877 
Thyroscyphus marginatus (Allman, 1877), new combination 
Fig. 8 


Obelia marginata Allman, 1877:9, pl. 6, figs. 1, 2. 

Campanularia insignis Fewkes, 1881:129. 

Not Campanularia marginata Bale, 1884:154, pl. 1, fig. 2. 

Lytoscyphus marginata.—Ritchie, 1909b:260. 

Lytoscyphus marginatus.—Billard, 1910:8. 

Campanularia marginata.—Nutting, 1915:44, pl. 6, figs. 5—7. 

Leptoscyphus marginatus.—Jaderholm, 1920:3. 

Cnidoscyphus marginatus.—Splettstosser, 1929:88, 125, figs. 83-88.—Calder, 1976: 
169.—Calder and Hester, 1978:91. 


Collection record.—Sta. B002, dredge, 12 Aug 1974. 

Description.—Specimen fragmentary, 2.8 cm high; hydrocaulus 0.8 mm wide, 
monosiphonic, straight basally, zigzag distally, divided by transverse nodes; peri- 
sarc thick. Each internode with a distal apophysis bearing a hydrotheca; every 
third internode with a second apophysis near the first; second apophysis large, 
supporting a branch; branches and hydrothecae given off alternately from op- 
posite sides of hydrocaulus. Branches nearly perpendicular to hydrocaulus, lack- 
ing secondary branches in present material; internodes zigzag, similar to those 
of hydrocaulus but more slender. Hydrothecae 1112—1264 um long from apoph- 
ysis to margin, cone-shaped; pedicel very short; base asymmetrical, with adcau- 


VOLUME 96, NUMBER 1 17 


line wall convex, abcauline wall nearly straight; diaphragm distinct, basal cham- 
ber small; margin entire, with a ring-like edge; aperture round, 562-655 um wide. 

Gonothecae absent. 

Remarks.—I follow Millard (1975) in regarding the genus Cnidoscyphus Splett- 
stosser, 1929, as a synonym of Thyroscyphus Allman, 1877. Obelia marginata 
Allman, 1877 from the Atlantic belongs to Thyroscyphus, as does Campanularia 
marginata Bale, 1884, from Australia (Bale 1915). A new name, Thyroscyphus 
balei, is proposed here for the junior secondary homonym 7. marginatus (Bale 
1884). Thyroscyphus marginatus is common to abundant in tropical and subtrop- 
ical waters of the western North Atlantic, apparently reaching the northern limit 
of its range off the coast of the Carolinas (Nutting 1915; Fraser 1944; Cain 1972). 
While it is frequent on hard banks off the coasts of Georgia and the Carolinas, 
only one fragmentary colony was collected during this study in estuarine areas 
of South Carolina. 

Known range.—Western Atlantic: North Carolina to Venezuela; Bermuda. 
Elsewhere: Tropical west coast of Africa. 


Family Plumulariidae L. Agassiz, 1862 
Subfamily Halopterinae Millard, 1962 
Monostaechas Allman, 1877 
Monostaechas quadridens (McCrady, 1859) 
Fig. 9 


Plumularia quadridens McCrady, 1859:199. 

Monostaechas dichotoma Aliman, 1877:37, pl. 22, figs. 1-5. 

Monostaechas quadridens.—Nutting, 1900:75, pl. 13, figs. 1-4.—Fraser, 1912: 
380, fig. 46; 1943:95; 1944:334, fig. 343.—Calder, 1976: 169.—Calder and Hester, 
1978:91. 

Monostaechas fisheri var. simplex Billard, 1913:16, fig. 7 (not Monostaechas 
fisheri Nutting, 1905). 

Monostaechas quadridens f. stechowi Leloup, 1935:2, figs. 2, 3. 


Collection record.—Sta. P006, trawl, 12 Jan 1976. 

Description.—Colonies reaching 8.5 cm high; hydrocaulus monosiphonic, 260 
ym in diameter, perisarc thick. Basal internodes very long, athecate, with nu- 
merous movable, 2 chambered nematothecae, most of which are missing in pres- 
ent specimens, giving rise via two curved apophyses to a pair of opposite upward- 
directed branches just below a hydrothecate segment, athecate and thecate seg- 
ments of hydrocaulus separated by an oblique hinge-joint. Branches resembling 
hydrocaulus and rebranched in like manner, the whole colony thus appearing to 
be dichotomously branched; some distal branches with a single branch-bearing 
apophysis below thecate segment. Hydrothecate segments up to 1 cm long, con- 
sisting of alternating hydrothecate and athecate internodes. Thecate internodes 
456-644 um long with an oblique hinge-joint proximally and a transverse node 
distally, each such internode with a hydrotheca, a movable median inferior ne- 
matotheca with a scoop-shaped aperture, a reduced median superior nematothe- 
ca, and a pair of movable trumpet-shaped lateral nematothecae on distinct apoph- 
yses adnate to hydrothecal wall. Athecate internodes 316-538 um long with a 
transverse node proximally and an oblique hinge-joint distally, with 1-2 movable 


18 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


nematothecae, each with a scoop-shaped aperture. Hydrotheca cup-shaped, 263— 
316 wm deep; main axis oblique to internode; abcauline wall slightly convex, 222— 
269 um long; free part of adcauline wall straight or slightly concave, 105-164 ~m 
long; margin entire, circular, aperture diameter 269-322 um. 

Gonothecae absent. 

Remarks.—McCrady’s (1859) type-material of M. quadridens from the Charles- 
ton Harbor area was found floating in the water, and specimens collected in an 
otter trawl from Port Royal Sound during this study were also all unattached. 
Attached hydroids of M. quadridens are abundant on ledges of the inner conti- 
nental shelf off South Carolina, and the species is probably carried into estuaries 
of the state by water currents. 

Known range.—Western Atlantic: Massachusetts to Brazil. Elsewhere: Cir- 
cumglobal, tropical and subtropical waters. 


Schizotricha Allman, 1883 
Schizotricha tenella (Verrill, 1874) 
Fig. 10 


Plumularia tenella Verrill, 1874:731. 

Schizotricha tenella.—Nutting, 1900:80, pl. 4, figs. 4, 5.—Fraser, 1912:383, fig. 
52.—Calder, 1976:169.—Calder and Hester, 1978:91. 

Halopteris tenella.—Vervoort, 1968:108. 


Collection records.—Sta. LRA7, dredge, 21 Apr 1976. Sta. LRA8, dredge, 21 
Apr 1976. Sta. MI07, dredge, 29 May 1975. Sta. MI09, dredge, 29 May 1975. Sta. 
MI12, dredge, 29 May 1975. Sta. MI13, dredge, 29 May 1975. Sta. MI15, dredge, 
29 May 1975. Sta. MI16, dredge, 29 May 1975. Sta. MI17, dredge, 29 May 1975. 
Sta. BBO, dredge, 11 Aug 1977. Sta. BBO2, dredge, 11 Aug 1977. Sta. SBO03, 
dredge, 22 Aug 1977, 28 Apr 1978, 25 July 1978. Sta. SB04, dredge, 22 Aug 1977, 
25 July 1978. Sta. SBO8, dredge, 22 Aug 1977, 25 May 1978, 25 July 1978. Sta. 
SB13, dredge, 22 Aug 1977, 28 Apr 1978, 25 July 1978. Sta. SB18, dredge, 22 
Aug 1977, 25 Apr 1978, 25 July 1978. Sta. BOO2, dredge, 15 Apr 1974, 22 Oct 
1974, 5 Jan 1976. Sta. BOO], dredge, 2 Apr 1976. Sta. E001, dredge, 2 Apr 1974, 
2 Aug 1974. Sta. E003, dredge, 2 Apr 1974, 2 Aug 1974, 3 Oct 1974. Sta. E008, 
dredge, 2 Oct 1974. Sta. DEOI, dredge, 13 June 1978. Sta. DEOS, dredge, 21 Nov 
1977, 15 Dec 1977, 13 June 1978, 17 Oct 1978. Sta. DE06, dredge, 21 Nov 1977, 
15 Dec 1977, 13 June 1978. Sta. DEO7, dredge, 21 Nov 1977, 17 Oct 1978. Sta. 
DE10, dredge, 21 Nov 1977, 13 June 1978, 17 Oct 1978. Sta. BRO8, dredge, 27 
Mar 1974, 8 Oct 1975. 

Description.—Colonies up to 3 cm high. Hydrocaulus monosiphonic, slightly 
zigzag, unbranched, basal region with hydrothecae, nematothecae, and hydro- 
cladia often missing, divided beyond basal region into alternating hydrothecate 
and athecate internodes. Internodes of variable length and diameter, usually long- 
er and thicker toward proximal end of hydrocaulus. Hydrothecate internodes with 
an oblique joint proximally and a transverse or slightly oblique node distally, the 
latter often indistinct; each thecate internode with one hydrotheca, one median 
inferior nematotheca, 2 lateral nematothecae, an occasional median superior ne- 
matotheca, and an apophysis near base of hydrotheca. Athecate internodes with 
a slightly oblique node proximally and an oblique joint distally, 1-4 nematothecae. 


VOLUME 96, NUMBER 1 19 


13 


12 


Figs. 9-13. Family Plumulariidae. 9, Monostaechas quadridens, hydrotheca, station P006; 10, 
Schizotricha tenella, hydrothecae and gonotheca, station BR08; 11, Plumularia floridana, hydrothe- 
cae and gonotheca, station BO01; 12, Aglaophenia trifida, hydrothecae, station CH00; 13, Macro- 
rhynchia philippina, hydrothecae, station BR08, scale bars = 500 um. 


Apophyses long, given off alternately from opposite sides of hydrocaulus, sup- 
porting unbranched or alternately branched hydrocladia. Hydrocladia up to 7 mm 
long, typically consisting of a series of 3 types of internodes; a very short athecate 
internode lacking nematothecae, nodes transverse at both ends; a longer athecate 
internode with 0-2 trumpet-shaped median nematothecae, proximal node trans- 
verse, distal node oblique; a long hydrothecate internode with a median inferior 
nematotheca, a hydrotheca, and a pair of trumpet-shaped lateral nematothecae, 
node oblique proximally, transverse distally. Nematothecae on both hydrocaulus 
and hydrocladia 2-chambered, movable. Hydrotheca cup-shaped, main axis oblique 
to hydroclade, abcauline wall nearly straight, 111-158 um long, free adcauline 
wall concave, length 70-94 wm, margin entire, opening circular, aperture diameter 
140-176 um. 


20 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Gonotheca cornucopia-shaped with 3 nematothecae near base, 562-690 um 
long, 263-316 wm wide, borne on a 2-segmented pedicel INSerene near base of 
hydrotheca on hydrocaulus and hydrocladia. 

Remarks.—Vannucci (1949) included this species in the synonymy of Halop- 
teris diaphana (Heller, 1868). I regard the two as separate species for the follow- 
ing reasons. The presence of three types of hydrocladial internodes in S. tenella 
immediately distinguishes it from H. diaphana, which has but two types. Hydro- 
cladia are usually branched in S. tenella and unbranched in H. diaphana. Finally, 
the largely temperate water S. tenella attains a much larger colony size than the 
tropical and subtropical H. diaphana. 

Schizotricha tenella is the most widespread plumularian hydroid in temperate 
estuaries of the east coast of the United States (Nutting 1900, 1901; Fraser 1912, 
1944; Calder 1971, 1976; Watling and Maurer 1972; Calder and Hester 1978). It 
is both euryhaline, penetrating up—estuary to a salinity of about !15%c, and eury- 
thermal, having been collected during this study over a temperature range from 
932°C. 

Known range.—Western Atlantic: Massachusetts to the Caribbean Sea. Else- 
where: Eastern Pacific, from Southern California to Panama. 


Subfamily Plumulariinae Agassiz, 1862 
Plumularia Lamarck, 1816 
Plumularia floridana Nutting, 1900 
Fig. 11 


Plumularia floridana Nutting, 1900:59, pl. 2, figs. 4, 5.—Fraser, 1912:381, fig. 
49.—Calder, 1976: 169.—Calder and Hester, 1978:91. 


Collection records.—Sta. NSO1, dredge, 6 Apr 1976. Sta. SB03, dredge, 25 July 
1978. Sta. SB04, dredge, 25 July 1978. Sta. SBO8, dredge, 25 July 1978. Sta. 
SB18, dredge, 22 Aug 1977. Sta. B002, dredge, 15 Apr 1974, 12 Aug 1974. Sta. 
BOO1, dredge, 16 Oct 1974, 13 Oct 1975, 2 Apr 1976. Sta. CHO0, dredge, 13 Oct 
1975. Sta. CHO2, dredge, 10 Oct 1973. Sta. JO02, trawl, 9 Oct 1973. Sta. E003, 
dredge, 8 Jan 1974, 2 Aug 1974, 3 Oct 1974. Sta. E005, dredge, 3 Oct 1974. Sta. 
E007, dredge, 2 Oct 1973. Sta. ACO1, Petersen grab, 6 June 1973. Sta. DEOI, 
dredge, 21 Oct 1977. Sta. DE04, dredge, 21 Nov 1977, 13 June 1978. Sta. DEOS, 
dredge, 13 June 1978, 17 Oct 1978. 

Description.—Hydrocaulus monosiphonic, straight, unbranched, reaching 2.5 
cm high, divided beyond basal region by distinct, transverse nodes; internodes 
293-480 um long, 70-105 wm wide at nodes, each with a distal apophysis and 2 
nematothecae, one beside apophysis and another proximal one on side opposite 
apophysis; apophyses given off alternately from opposite sides of hydrocaulus, 
giving rise to unbranched hydrocladia with 1-5 hydrothecae; hydrocladia up to 2 
mm long, nodes alternately transverse and oblique, hydroclade with 1-3 short 
basal internodes lacking hydrothecae and nematothecae, remaining internodes 
alternately hydrothecate and athecate, internodes with 2 internodal septa, one 
near each extremity, often indistinct; thecate internodes 257-351 wm long with a 
median inferior nematotheca, a hydrotheca, and a pair of lateral nematothecae, 
lateral nematothecae not reaching hydrothecal margin; athecate internodes 94— 
257 wm long, usually with one median nematotheca, all nematothecae movable, 


VOLUME 96, NUMBER 1 1 


2-chambered, cone-shaped; hydrotheca cup-shaped, main axis oblique to hydro- 
clade, abcauline wall 129-164 um long, length adcauline wall free 70-105 um, 
margin entire, aperture diameter 117-164 um. 

Female gonotheca oval with thin perisarc, 211-281 wm long, 99-123 um wide 
when fully developed, borne on slender pedicels from anterior side of apophysis 
base, each containing one ovum or developing planula, additional ova visible in 
coenosarc of hydrocaulus, planula escaping from rupture at distal end of gono- 
theca. 

Remarks.—This species closely resembles descriptions of Plumularia alicia 
Torrey, 1902 from California and Oregon, USA (Torrey 1902; Fraser 1937), Plu- 
mularia sp. from Queensland, Australia (Pennycuik 1959), and P. pennycuikae 
Millard and Bouillon, 1973 from the Seychelles (Millard and Bouillon 1973), Bonin 
Islands (Hirohito 1974), and Mozambique (Millard 1975). Plumularia floridana 
differs from these species chiefly in having one instead of two nematothecae in 
the axil between hydrocaulus and apophysis. Plumularia indica from India (Mam- 
men 1965b) is similar, and like P. floridana has a single axillary nematotheca. If 
not conspecific, these hydroids must all be very closely related. 

Known range.—Western Atlantic: North Carolina to Brazil; Bermuda; north- 
ward to Massachusetts on floating Sargassum. Elsewhere: Eastern Pacific, from 
Southern California to Ecuador. 


Subfamily Aglaopheniinae Stechow, 1911 
Aglaophenia Lamouroux, 1812 
Aglaophenia trifida L. Agassiz, 1862 
Fig. 12 


Aglaophenia cristata.—McCrady, 1859:202 [not Plumularia cristata Lamarck, 
1816 = Aglaophenia pluma (Linnaeus, 1758)]. 

Aglaophenia trifida L. Agassiz, 1862:358.—Calder, 1976:169.—Calder and Hes- 
ter, 1978:91. 

Aglaophenia rigida Allman, 1877:43, pl. 25, figs. 5—9.—Jaderholm, 1896:17.— 
Fraser, 1912:378, fig. 44. 


Collection records.—Sta. YB02, dredge, 6 Jan 1977. Sta. NSOO, trawl, 17 Dec 
1974. Sta. NBO2, trawl, 19 Dec 1974. Sta. CHO0, trawl, 22 May 1974. 

Description.—Colonies growing in tangled masses reaching 20 cm high. Hy- 
drocaulus monosiphonic, wiry, slender, reaching 0.5 mm wide basally, occasion- 
ally unbranched but usually with long, widely-separated branches. Branches giv- 
en off from front of hydrocaulus either singly or in pairs, frequently rebranched 
in like manner; paired branches when present arising from contiguous internodes. 
Hydrocaulus and branches divided by oblique nodes into short internodes mea- 
suring 300-560 um long, each internode with one apophysis and 3 nematothecae, 
one proximal to, one on, and one in the axil of each apophysis, apophyses given 
off alternately from fronto-lateral surface of internodes. Hydrocladia reaching 13 
mm long, nearly straight, absent only at base of hydrocaulus and some older 
branches, divided at regular intervals by slightly oblique nodes; internodes 281— 
351 um long, 94-164 wm wide at nodes, each with one hydrotheca, one median 
inferior nematotheca, one pair of lateral nematothecae, and 2 internodal septa, 
one at base of intrathecal septum, the other at base of lateral nematothecae. 


i) 
N 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Hydrothecae cone-shaped, 269-328 um long, 164-187 um wide at aperture, oc- 
cupying most of internode, orifice of one hydrotheca adjacent to base of next, 
adcauline wall convex, distal half of abcauline wall concave with thick perisarc, 
plane of orifice sloping. Margin with one median and 4 pairs of lateral teeth, teeth 
rounded, short intrathecal septum present basally. Nematothecae tubular, one- 
chambered, immovable; median nematotheca adnate to hydrotheca for much of 
its length, reaching halfway to hydrothecal margin, length free 23-47 um; lateral 
nematothecae adnate to hydrotheca, reaching nearly to orifice. 

Corbulae pod-shaped, 4 mm long, 770 wm wide, each occurring in place of a 
distal hydrocladium, borne on short pedicels consisting of an internode with one 
hydrotheca; either side of corbula with 11-13 gonohydrocladia, each with a row 
of about 6 nematophores; interior of corbula with oval gonothecae. 

Remarks.—Two distinct colony forms of this species were found in estuaries 
of South Carolina. Hydroids from several shallow subtidal and intertidal areas 
(stations YBO2, NS00, NBO2), appeared to be dichotomously branched because 
the long branches arose singly from the hydrocaulus. A different colony form 
was evident in specimens collected from deeper water at station CH00 near the 
Charleston Harbor entrance. These hydroids had long branches arising in pairs 
from the hydrocaulus, their hydrocladia were longer, and the diameters of their 
hydrocauli were greater. Nevertheless, hydrothecae of the two forms were in- 
distinguishable and the variation in colony form is believed to be environmentally 
induced. 

Specimens of this species from Sullivans Island and Charleston, South Caro- 
lina, were identified by McCrady (1859) as Aglaophenia cristata. However, he 
believed that it was probably distinct from the species described from Europe by 
Lamarck (1816) as Plumularia cristata [=Aglaophenia pluma (Linnaeus, 1758)]. 
Agassiz (1862) provided a new name, A. trifida, for this hydroid but did not 
describe or illustrate it. Despite Fraser’s (1944) apparent views to the contrary, 
Agassiz should be recognized as the author of the species because he provided 
the necessary “‘indication’’ required under Article 16 (ICZN). 

There is nothing in the original description of Aglaophenia rigida by Allman 
(1877), or in subsequent descriptions of that species by later authors, that can be 
used to distinguish it from A. trifida. Although Nutting (1900) and Fraser (1944) 
reported differences in the number of marginal teeth between the two species, 
with eight in A. rigida and nine in A. trifida, such differences are nonexistent. 
The types of A. rigida (MCZ 2109) have nine marginal teeth. Accordingly, the 
Species are regarded here as synonyms, with the name A. trifida L. Agassiz, 1862 
having priority over A. rigida Allman, 1877. 

Aglaophenia trifida bears considerable resemblance to descriptions of the cos- 
mopolitan A. pluma (Linnaeus, 1758). Nutting (1900) compared specimens of A. 
trifida with material of A. pluma from Plymouth, England, and concluded that 
the two were different species. He observed that the abcauline wall of the hy- 
drotheca distal to the median inferior nematotheca is less concave in A. trifida 
and that the corbulae of this species were much longer and more slender. For 
the present I prefer to treat A. trifida as a distinct species, but it may eventually 
be shown to be conspecific with A. pluma. Records of A. pluma from the western 
North Atlantic as A. dichotoma (Leloup 1937; Fraser 1944) and A. pluma pluma 


VOLUME 96, NUMBER | 23 


(van Gemerden-Hoogeveen 1965) are probably based on hydroids identical with 
those identified here as A. trifida. 
Known range.—Western Atlantic: North Carolina to the Caribbean Sea. 


Macrorhynchia Kirchenpauer, 1872 
Macrorhynchia philippina Kirchenpauer, 1872 
Fig. 13 


Aglaophenia (Macrorhynchia) philippina Kirchenpauer, 1872:29, 45, pls. 1, 2, 7, 
ne 2b: 

Aglaophenia urens.—Bale, 1884:155, pl. 14, fig. 6, pl. 17, fig. 9 (not Aglaophenia 
urens Kirchenpauer, 1872). 

Lytocarpus phillipinus.—Bale, 1888:786, pl. 21, figs. 5—7 (lapsus). 

Lytocarpus philippinus.—Marktanner-Turneretscher, 1890:274, pl. 6, fig. 16.— 
Fraser, 1912:379, fig. 45.—Calder, 1976:169.—Calder and Hester, 1978:91. 

Lytocarpia philippina.—Stechow, 1919:132, fig. Z1. 

Macrorhynchia philippina.—Stechow, 1923:241. 


Collection records.—Sta. B003, dredge, 12 Aug 1974. Sta. BO02, dredge, 13 
Oct 1975. Sta. CIO1, trawl, 5 Oct 1976. Sta. BRO8, dredge, 8 Oct 1975. Sta. P006, 
trawl, 7 Aug 1974. 

Description.—Irregularly, pinnately, or twice-pinnately branched colonies 
reaching 10 cm high. Hydrocaulus and branches curved gradually backward, 
polysiphonic except at distal ends, bearing hydrocladia except when these are 
broken off proximally; nodes indistinct; apophyses short, given off alternately 
from front of anterior stolon only, each apophysis bearing one rudimentary ne- 
matotheca and one axillary nematotheca, an additional nematotheca on stolon 
between adjacent apophyses. Hydrocladia up to 7 mm long, curved outward, 
nodes straight; internodes 257-374 um long, 75-100 um wide at nodes, each with 
one hydrotheca, one median inferior nematotheca, one pair of lateral nematothe- 
cae, and 2 septa. Hydrothecae boot-shaped, 257-316 um long, 140-187 wm wide 
at aperture; abcauline wall with a distinct inward-projecting intrathecal septum, 
adcauline wall with a short intrathecal septum basally; margin with low, rounded 
lateral teeth and an acute median tooth. Nematothecae tubular, one-chambered, 
immovable; median inferior nematotheca long, adnate to hydrotheca proximally, 
reaching beyond margin of hydrotheca distally, length free 82-140 um, internal 
aperture diameter 11-19 um; lateral nematothecae projecting a short distance 
beyond hydrothecal margin distally, otherwise adnate to hydrotheca. 

Phylactocarpia up to 2.5 mm long, occurring in place of hydrocladia. Basal 
internode of phylactocarp with a hydrotheca, a median inferior nematotheca, a 
pair of lateral nematothecae, and 2 internodal septa; terminal internode also hy- 
drothecate, or with nematothecae only; intermediate internodes modified, one or 
more bearing gonothecae, nematothecae well-developed. Gonothecae disc-shaped, 
1217-1287 wm wide, 1193-1463 wm high, 550-667 um thick; 1-3 per phylactocarp. 

Remarks.—Following authors such as Stechow (1923), Vannucci Mendes (1946), 
Mammen (1965b) and Gravier (1970), Kirchenpauer’s (1872) subgenus Macro- 
rhynchia is recognized as a distinct genus, and Lytocarpus Allman, 1883 is re- 
garded as a synonym. 


24 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


According to the literature (Gravely 1927; Halstead 1965), this hydroid is ca- 
pable of stinging humans. 

Known range.—Western Atlantic: North Carolina to Brazil; Bermuda. Else- 
where: Circumglobal, tropical and subtropical waters. 


Acknowledgments 


Materials for this study were collected while I was a staff member of the Marine 
Resources Research Institute, Charleston, South Carolina. Field work was sup- 
ported by funds from the Coastal Plains Regional Commission (Contract No. 
10340031), the U.S. Army Corps of Engineers (Contract Nos. DACW60-75-C- 
0016, DACW60-76-C-0017, DACW60-77-C-0013), and the South Carolina Coastal 
Council. Thanks are due to Magdalene Maclin, Billy Boothe, David Knott, and 
Dr. Robert Van Dolah for assistance in the field. I am also indebted to Dr. K. P. 
Sebens of the Museum of Comparative Zoology at Harvard University and Dr. 
F. M. Bayer of the National Museum of Natural History, Smithsonian Institution, 
for the loan of specimens. Mr. A. Sanders kindly permitted access to invertebrate 
collections at the Charleston Museum in a search for any hydrozoan materials 
collected by John McCrady. Finally, I thank an anonymous reviewer for con- 
structive criticisms. 


Literature Cited 


Agassiz, L. 1862. Contributions to the natural history of the United States of America. Volume 
4.—Little, Brown and Company, Boston, 380 pp. 

Alder, J. 1856. A notice of some new genera and species of British hydroid zoophytes.—Annals 
and Magazine of Natural History (2)18:353-362. 

Allman, G. J. 1877. Report on the Hydroida collected during the exploration of the Gulf Stream by 
L. F. de Pourtalés, assistant United States Coast Survey.—Memoirs of the Museum of Com- 
parative Zoology at Harvard College 5(2):1—66. 

—. 1883. Report on the Hydroida dredged by H.M.S. Challenger during the years 1873-1876. 
Part I. Plumularidae.—Report on the Scientific Results of the Voyage of H.M.S. Challenger 
during the Years 1873-1876, Zoology 7:1—54. 

——. 1885. Description of Australian, Cape and other Hydroida, mostly new, from the collection 
of Miss H. Gatty.—Journal of the Linnaean Society 19:132-161. 

——. 1888. Report on the Hydroida dredged by H.M.S. Challenger during the years 1873-1876. 
Part If. The Tubularinae, Corymorphinae, Campanularinae, Sertularinae, and Thala- 
mophora.—Report on the Scientific Results of the Voyage of H.M.S. Challenger during the 
Years 1873-1876, Zoology 23:1—90. 

Bale, W. M. 1884. Catalogue of the Australian hydroid zoophytes.—Australian Museum, Sydney, 

198 pp. 

1888. On some new and rare Hydroida in the Australian Museum collection.—Proceedings 
of the Linnaean Society of New South Wales 3:745-799. 

1913. Further notes on Australian hydroids. IIl.—Proceedings of the Royal Society of Vic- 
toria (new series) 26:114—147. 

1915. Report on the Hydroida collected in the Great Australian Bight and other localities. 
Part III.—Zoological Results of the Fishing Experiments carried out by F.I.S. ““Endeavour”’ 
3:241-336. 

Bennitt, R. 1922. Additions to the hydroid fauna of the Bermudas.—Proceedings of the American 
Academy of Arts and Sciences 57:241-259. 

Billard, A. 1910. Revision d’une partie des hydroides du British Museum.—Annales Sciences Na- 
turelles, Zoologie 11:1—67. 

——. 1912. Hydroides de Roscoff.—Archives de Zoologie Expérimentale et Générale 51:459- 
478. 


VOLUME 96, NUMBER 1 25 


——. 1913. Les hydroides de l’expédition du Siboga. I. Plumulariidae.—Siboga-Expeditie, Mon- 

ographie 7a:1—115. 

1919. Note sur quelques espéces nouvelles de Sertularella de l’expédition du ‘‘Siboga.’’— 
Archives de Zoologie Expérimentale et Générale 58:18—23. 

1924. Note sur quelques espéces la plupart nouvelles de synthecides et de sertularides du 
“‘Siboga.’’—Bulletin de la Société Zoologique de France 49:646—652. 

1925. Les hydroides de l’Expédition du Siboga. II. Synthecidae et Sertularidae.—Siboga- 
Expeditie, Monographie 7b: 115—232. 

Blanco, O. M. 1968. Nueva contribucion al conocimiento de la fauna marina hidroide.—Revista del 
Museo de la Plata (Nueva Serie), Seccion Zoologia 10:195—224. 

Brooks, W. K. 1886. The life-history of the Hydromedusae: A discussion of the origin of the 
medusae, and of the significance of metagenesis.—Memoirs of the Boston Society of Natural 
History 3:359-430. 

Cain, T. D. 1972. Additional epifauna of a reef off North Carolina.—Journal of the Elisha Mitchell 
Scientific Society 88:79-82. 

Calder, D. R. 1971. Hydroids and Hydromedusae of southern Chesapeake Bay.—Virginia Institute 

of Marine Science Special Papers in Marine Science 1, 125 pp. 

1976. The zonation of hydroids along salinity gradients in South Carolina estuaries.—Jn G. 
O. Mackie, ed., Coelenterate Ecology and Behavior, Plenum Press, New York, pp. 165-174. 
, and B. S. Hester. 1978. Phylum Cnidaria.—In R. G. Zingmark, ed., A Checklist of the 

Biota of the Coastal Zone of South Carolina, University of South Carolina Press, Columbia, 
pp. 87-93. 

Clarke, S. F. 1879. Report on the Hydroida collected during the exploration of the Gulf Stream 
and Gulf of Mexico by Alexander Agassiz, 1877—1878.—Bulletin of the Museum of Compar- 
ative Zoology at Harvard College 5:239-252. 

Cornelius, P. F. S. 1979. A revision of the species of Sertulariidae (Coelenterata: Hydroida) re- 
corded from Britain and nearby seas.—Bulletin of the British Museum (Natural History), 
Zoology Series 34:243-321. 

Coughtrey, M. 1876. Critical notes on the New Zealand Hydroida.—Transactions and Proceedings 
of the New Zealand Institute 8:298—302. 

Dean, T. A., and V. J. Bellis. 1975. Seasonal and spatial distribution of epifauna in the Pamlico 
River estuary, North Carolina.—Journal of the Elisha Mitchell Scientific Society 91:1—12. 

Ellis, J., and D. Solander. 1786. The natural history of many curious and uncommon zoophytes, 
collected from various parts of the globe by the late J. Ellis. . . . Systematically arranged and 
described by the late Daniel Solander. . . . London, 208 pp. 

Fewkes, J. W. 1881. Reports on the results of dredgings, under the supervision of Alexander 
Agassiz, in the Caribbean Sea, in 1878, 1879, and along the Atlantic coast of the United States 
during the summer of 1880, by the U.S. Coast Survey steamer “‘Blake.’’ XI. Report on the 
Acalephae.—Bulletin of the Museum of Comparative Zoology at Harvard College 8:127-140. 

Fleming, J. 1828. A History of British Animals. Bell and Bradfute, Edinburgh, 565 pp. 

Fraser, C. M. 1912. Some hydroids of Beaufort, North Carolina.—Bulletin of the United States 

Bureau of Fisheries 30:339-387. 

1913. Hydroids from Nova Scotia.—Victoria Memorial Museum Bulletin Number 1:157— 
186. 

1937. Hydroids of the Pacific coast of Canada and the United States.—University of Toronto 
Press, Toronto, 207 pp. 

———. 1943. Distribution records of some hydroids in the collection of the Museum of Comparative 
Zoology at Harvard College, with description of new genera and new species.—Proceedings 
of the New England Zoological Club 22:75-98. 

——. 1944. Hydroids of the Atlantic Coast of North America.—University of Toronto Press, 

Toronto, 451 pp. 

1948. Hydroids of the Allan Hancock Pacific expeditions since March, 1938.—Allan Han- 
cock Pacific Expeditions 4:179-335. 

Garcia, P., A. Aguirre, and D. Gonzalez. 1980. Contribucion al conocimiento de los hidrozoos de 
las costas Espanolas. Parte III. “‘Sertulariidae.’’—Boletin del Instituto Espanol de Oceano- 
grafia 6:5—67. 

Gemerden-Hoogeveen, G. C.H. van. 1965. Hydroids of the Caribbean: Sertulariidae, Plumulanidae 
and Aglaophentidae.—Studies on the Fauna of Curagao and other Caribbean Islands 22: 1-87. 


26 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Gravely, F. H. 1927. The littoral fauna of Krusadai Island in the Gulf of Manaar. Hydrozoa.— 
Bulletin of the Madras Government Museum (New Series, Natural History) 1:7—20. 

Gravier, N. 1970. Libération de médusoides par Macrorhynchia philippina Kirchenpauer, 1872 
(Hydroida, Plumulariidae).—Recueil des Travaux de la Station Marine d’ Endoume (Fascicule 
Hors Série), Supplement Numereau 10:253—257. 

Gray, J. E. 1848. List of the specimens of British animals in the collection of the British Museum. 
Part 1. Centroniae or radiated animals. London. 

Halstead, B. W. 1965. Poisonous and venomous marine animals of the world. J. Invertebrates.— 
United States Government Printing Office, Washington, D.C., 994 pp. 

Hargitt, C. W. 1908. Notes on a few coelenterates of Woods Hole.—Biological Bulletin 14:95—-120. 

Hartlaub, C. 1905. Die Hydroiden der magalhaenischen Region und chilenischen Kuste.—Zoolo- 
gische Jahrbticher, Supplement 6:497-714. 

Hassall, A. H. 1848. Definitions of three new British zoophytes.—Zoologist 6:2223. 

Heller, C. 1868. Die Zoophyten und Echinodermen des Adriatischen Meeres.—Verhandlungen der 
Zoologisch-Botanischen Gesellschaft in Wien 18:1-88. 

Hincks, T. 1868. A history of the British hydroid zoophytes, Vol. I.—John van Voorst, London, 
338 pp. 

Hirohito, Emperor of Japan. 1969. Some hydroids of the Amakusa Islands.—Biological Laboratory, 
Imperial Household, Tokyo, 32 pp. 

—. 1974. Some hydroids of the Bonin Islands.—Biological Laboratory, Imperial Household, 
Tokyo, 55 pp. 

Howard, J. D., and R. W. Frey. 1975. Estuaries of the Georgia coast, U.S.A.: Sedimentology and 
biology. I. Introduction.—Senckenbergiana Maritima 7:1—31. 

Jaderholm, E. 1896. Ueber aussereuropaische Hydroiden des Zoologischen Museums der Univer- 

sitat Upsala.—Bihang till Svenska Vetenskapsakademiens Handlingar 21 IV(6):1—20. 

1903. Aussereuropaische Hydroiden im Swedischen Reichsmuseum.—Arkiv for Zoologi 1: 


259-312. 
—. 1920. Onsome exotic hydroids in the Swedish Zoological State Museum.—Arkiv for Zoologi 
13(3): 1-11. 


Jarvis, F. E. 1922. The hydroids from the Chagos, Seychelles, and other islands and from the coasts 
of British East Africa and Zanzibar.—Transactions of the Linnaean Society of London (Zo- 
ology) 18:331—360. 

Karlson, R. 1978. Predation and space utilization patterns in a marine epifaunal community.— 
Journal of Experimental Marine Biology and Ecology 31:225—239. 

Kirchenpauer, G. H. 1864. Neue Sertulariden aus verschiedenen Hamburgischen Sammlungen, 
nebst allgemeinen Bemerkungen tiber Lamouroux’s Gattung Dynamena.—Verhandlungen der 
K. Leopoldinisch-Carolinischen Deutschen Akademie der Naturforscher 31(3):1—16. 

——. 1872. Ueber die Hydroidenfamilie Plumularidae, einzelne Gruppen derselben und ihre 
Fruchtbehalter. I. Aglaophenia Lx.—Abhandlungen aus dem Gebiete der Naturwissenschaf- 
ten, Hamburg 5:1—52. 

Kuhn, A. 1913. Entwicklungsgeschichte und Verwandtschaftsbeziehungen der Hydrozoen. I. Teil. 
Die Hydroiden.—Ergebnisse und Fortschritte der Zoologie 4:1—284. 

Lamarck, J. P. B. A. de M. 1816. Histoire naturelle des animaux sans vertebres. Ed. 2. Paris, 
568 pp. 

Lamouroux, J. V. F. 1812. Extrait d'un mémoire sur la classification des polypiers coralligenes non 
entierement pierreux.—Nouveau Bulletin des Sciences pour la Société Philomatique de Paris 


3:181-188. 
—. 1816. Histoire des polypiers coralligénes flexibles, vulgairement nommeés zoophytes. Caen, 
560 pp. 


—. 1821. Exposition méthodique des genres de ]’ordre des polypiers, avec leur description et 
celle des principales espéces. Paris, 115 pp. 

Leloup, E. 1935. Hydropolypes calyptoblastiques et siphonophores récoltés au cours de la croisiére 
(1934-35) du navire-école Belge “‘Mercator’’.—Bulletin du Musée Royal d’Histoire Naturelle 
de Belgique 11:1-6. 

—. 1937. Résultats scientifiques des croisiéres du navire-école Belge ‘‘Mercator’’. Vol. I. Hy- 
droida.—Mémoires du Musée Royal d’ Histoire Naturelle de Belgique 2, 9:91—121. 

Linnaeus, C. 1758. Systema Naturae per Regna Tria Naturae, secundem Classes, Ordines, Genera, 
Species, cum characteribus, differentiis, Synonymis, Locis, Editio Decima, Reformata. Hol- 
miae, 824 pp. 


VOLUME 96, NUMBER 1 2H 


Mammen, T. A. 1965a. On a collection of hydroids from South India. II. Suborder Thecata (ex- 
cluding Family Plumulartidae).—Journal of the Marine Biological Association of India 7:1—57. 

——. 1965b. On a collection of hydroids from South India. III. Family Plumulariidae.—Journal 
of the Marine Biological Association of India 7:291-324. 

Marktanner-Turneretscher, G. 1890. Die Hydroiden des K.K. naturhistorischen Hofmuseums.— 
Annalen des Naturhistorischen Museums in Wien 5:195—286. 

Mayer, A.G. 1910a. Medusae of the world. I. The Hydromedusae.—Carnegie Institution of Wash- 
ington Publication 109: 1—230. 

———. 1910b. Medusae of the world. Il. The Hydromedusae.—Carnegie Institution of Washington 
Publication 109:231—498. 

McCrady, J. 1859. Gymnopthalmata of Charleston Harbor.—Proceedings of the Elliott Society of 
Natural History 1:103—221. 

McDougall, K. D. 1943. Sessile marine invertebrates of Beaufort, North Carolina.—Ecological 
Monographs 13:321-374. 

Millard, N. A. H. 1962. The Hydrozoa of the south and west coasts of South Africa. Part I. The 
Plumulariidae.—Annals of the South African Museum 46:261-319. 

——. 1964. The Hydrozoa of the south and west coasts of South Africa. Part II. The Lafoeidae, 
Synthecidae and Sertulariidae.—Annals of the South African Museum 48:1—56. 

—. 1975. Monograph on the Hydroida of southern Africa.—Annals of the South African Mu- 

seum 68:1—513. 

, and J. Bouillon. 1973. MHydroids from the Seychelles (Coelenterata)—Musée Royal de 

| Afrique Centrale, Annales, Serie IN-8, Sciences Zoologiques 206:1—106. 

, and 1974. A collection of hydroids from Mogambique, East Africa.—Annals of the 

South African Museum 65:1—40. 

Nutting, C.C. 1900. American hydroids. Part I. The Plumularidae.—Smithsonian Institution, United 
States National Museum Special Bulletin 4(1):1—285. 

——. 1901. The hydroids of the Woods Hole region.—Bulletin of the United States Bureau of 
Fisheries 19:325—386. 

—. 1904. American hydroids. Part II. The Sertularidae.—Smithsonian Institution, United States 
National Museum Special Bulletin 4(2):1—325. 

——. 1905. Hydroids of the Hawaiian Islands collected by the steamer Albatross in 1902.— 
Bulletin of the United States Fish Commission 23:93 1-959. 

——. 1915. American hydroids. Section III. The Campanularidae and Bonneviellidae.—Smith- 
sonian Institution, United States National Museum Special Bulletin 4(3): 1-126. 

Pearse, A. S. 1936. Estuarine animals at Beaufort, North Carolina.—Journal of the Elisha Mitchell 

Scientific Society 52:174—222. 

, and L. G. Williams. 1951. The biota of the reefs off the Carolinas.—Journal of the Elisha 

Mitchell Scientific Society 67:133-161. 

Pennycuik, P. R. 1959. Faunistic records from Queensland. Part V.—Marine and brackish water 
hydroids.—Papers from the Department of Zoology, University of Queensland 1:141—210. 

Pictet, C. 1893. Etude sur les hydraires de la Baie d’ Amboine.—Revue Suisse de Zoologie 1:1—64. 

Ralph, P. M. 1961. New Zealand thecate hydroids. Part I[I—Family Sertulariidae.—Transactions 
of the Royal Society of New Zealand 88:749-838. 

Ritchie, J. 1909a. Supplementary report on the hydroids of the Scottish National Antarctic Expe- 
dition.—Transactions of the Royal Society of Edinburgh 47:65-101. 

——. 1909b. Two unrecorded Challenger hydroids from the Bermudas with a note on the syn- 
onymy of Campanularia insignis.—Zoologist 13:260-263. 

Sandifer, P. A., T. I. J. Smith, and D. R. Calder. 1974. Hydrozoans as pests in closed-system 
culture of larval decapod crustaceans.—Aquaculture 4:55—59. 

Splettstosser, W. 1929. Beitrage zur Kenntnis der Sertulariiden. Thyroscyphus Allm., Cnidoscyphus 
nov. gen., Parascyphus Ritchie.—Zoologische Jahrbticher, Systematik, Okologie und Geogra- 
phie der Tiere 58:1—134. 

Stechow, E. 1911. Uber Hydroiden der Deutschen Tiefsee-Expedition. Ein neues Genus thekater 
Hydroiden.—Zoologischer Anzeiger 37: 193-197. 

——. 1919. Zur Kenntnis der Hydroidenfauna des Mittelmeeres, Amerikas und anderer Gebiete, 
nebst Angaben Uber einige Kirchenpauer’sche Typen von Plumulariden.—Zoologische Jahr- 
biicher, Systematik, Okologie und Geographie der Tiere 42: 1-172. 

———. 1921. Neue Genera und Species von Hydrozoen und anderen Evertebraten.—Archiv fur 
Naturgeschichte 87:248—265. 


28 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


——. 1923. Zur Kenntnis der Hydroidenfauna des Mittelmeeres, Amerikas und anderer Ge- 

biete.—Zoologische Jahrbiicher, Systematik, Okologie und Geographie der Tiere 47:29-270. 

, and H. C. Miller. 1923. Hydroiden der Aru-Inseln.—Senckenbergische Naturforschende 

Gesellschaft 35:459-478. 

Stephenson, T. A., and A. Stephenson. 1952. Life between tide-marks in North America. II. North- 
ern Florida and the Carolinas.—Journal of Ecology 40:1—49. 

Sutherland, J. P. 1977. Effect of Schizoporella (Ectoprocta) removal on the fouling community at 

Beaufort, North Carolina, USA.—Jn B. C. Coull, ed., Ecology of marine benthos. University 

of South Carolina Press, Columbia, pp. 155-176. 

1981. The fouling community at Beaufort, North Carolina: A study in stability —American 
Naturalist 118:499-519. 

, and R. H. Karlson. 1977. Development and stability of the fouling community at Beaufort, 

North Carolina.—Ecological Monographs 47:425-446. 

Thompson, D’A. W. 1879. On some new and rare hydroid zoophytes (Sertulariidae and Thuiariidae) 
from Australia and New Zealand.—Annals and Magazine of Natural History (5)3:97—114. 

Thornely, L. R. 1904. Report on the Hydroida collected by Professor Herdman, at Ceylon, in 
1902.—Report to the Government of Ceylon on the Pearl Oyster Fisheries of the Gulf of Manaar 
II, Supplementary Report 8:107—126. 

Torrey, H. B. 1902. The Hydroida of the Pacific coast of North America, with especial reference 
to the species in the collection of the University of California.—University of California Pub- 
lications, Zoology 1:1-104. 

Totton, A. K. 1930. Coelenterata.—Part V.—Hydroida.—Bnitish Antarctic (“Terra Nova’’) Ex- 
pedition, 1910, Natural History Reports, Zoology 5(5):131—252. 

Vannucci, M. 1949. Hydrozoa do Brasil.—Boletim da Faculdade de Filosofia, Ciéncias e Letras, 
Universidade de Sao Paulo, Zoologia 14:219-265. 

Vannucci Mendes, M. 1946. Hydroida Thecaphora do Brasil.—Arquivos de Zoologia do Estado de 
Sao Paulo 4:535-597. 

Verrill, A. E. 1872. On Radiata from the coast of North Carolina.—American Journal of Science 

(3)3:432-438. 

1874. Report upon the invertebrate animals of Vineyard Sound and the adjacent waters, 
with an account of the physical features of the region.—Report of the Commissioner of Fish- 
eries for 1871—1872:295-747. 

—. 1900. Additions to the Anthozoa and Hydrozoa of the Bermudas.—Transactions of the 
Connecticut Academy of Arts and Sciences 10:551—572. 

—. 1907. Characteristic life of the Bermuda coral reefs.—Transactions of the Connecticut 
Academy of Arts and Sciences 12:204—319. 

Versluys, J., Jr. 1899. Hydraires calyptoblastes recueillis dans la Mer des Antilles pendant lune 
des croisi¢res accomplies par le Comte R. de Dalmas sur son yacht Chazalie—Mémoires de 
la Société Zoologique de France 12:29-58. 

Vervoort, W. 1968. Report on a collection of Hydroida from the Caribbean region, including an 

annotated checklist of Caribbean hydroids.—Zoologische Verhandelingen 92:1—124. 

1972. Hydroids from the Theta, Vema, and Yelcho cruises of the Lamont-Doherty Geo- 
logical Observatory.—Zoologische Verhandelingen 120:1—247. 

, and P. Vasseur. 1977. Hydroids from French Polynesia with notes on distribution and 

ecology.—Zoologische Verhandelingen 159:1—98. 

Watling, L., and D. Maurer. 1972. Shallow water hydroids of the Delaware Bay region.—Journal 
of Natural History 6:643-649. 

Watson, J. E. 1973. Pearson Island Expedition 1969-9. Hydroids.—Transactions of the Royal So- 
ciety of South Australia 97:153—200. 

Wells, H. W. 1961. The fauna of oyster beds, with special reference to the salinity factor.—Eco- 
logical Monographs 31:239-266. 

———,, M. J. Wells, and I. E. Gray. 1964. The calico scallop community in North Carolina.— 
Bulletin of Marine Science of the Gulf and Caribbean 14:561—593. 


Department of Invertebrate Zoology, Royal Ontario Museum, 100 Queen’s 
Park, Toronto, Ontario MSS 2C6, Canada. 


PROC. BIOL. SOC. WASH. 
96(1), 1983, pp. 29-37 


CREEDIA ALLENI AND CREEDIA PARTIMSQUAMIGERA 
(PERCIFORMES: CREEDIIDAE), TWO NEW MARINE 
FISH SPECIES FROM AUSTRALIA, WITH 
NOTES ON OTHER AUSTRALIAN 
CREEDIIDS 


Joseph S. Nelson 


Abstract.—Two new species of trachinoid fishes, Creedia alleni and C. par- 
timsquamigera, are described. Creedia alleni, known from three specimens col- 
lected along coastal Western Australia, is distinguished from all other creediids 
in having only three soft rays, in addition to a spine, in each pelvic fin. Creedia 
partimsquamigera, known from nine specimens from New South Wales, is similar 
to C. haswelli (Ramsay) in fin ray counts but differs in lacking scales on the 
anterior portion of the body except along the lateral line and before the dorsal 
fin, and differs further from the other two species of Creedia in having a blunter 
snout and a blunt maxilla extending well behind the eye. Creedia partimsqua- 
migera and C. haswelli exhibit sexual dimorphism, with males having longer 
pectoral and pelvic fins than females. Notes are given on the three other creediids 
from Australia, namely, C. haswelli, Limnichthys fasciatus Waite, and L. don- 
aldsoni Schultz (newly found in Australia). 


The family Creediidae (including Limnichthyidae) currently contains 12 de- 
scribed species in seven genera (Nelson 1979). Herein I describe two new species, 
one from near Perth, Western Australia, and the other from Sydney, New South 
Wales. New material of Limnichthys fasciatus from Western Australia and of L. 
donaldsoni and Creedia haswelli from eastern Australia is reported. 


Methods 


Measurements were made to the nearest 0.1 mm with needle-point dial calipers 
and are expressed as thousandths (%c) of standard length (SL). Radiographs were 
made of the specimens of the two new species. Abbreviations refer to the follow- 
ing museums: AMNH, American Museum of Natural History, New York; AMS, 
Australian Museum, Sydney; NMV, National Museum of Victoria, Melbourne; 
QVM, Queen Victoria Museum, Launceston; TFDA, Tasmanian Fisheries De- 
velopment Authority, Hobart; UAMZ, University of Alberta Museum of Zool- 
ogy, Edmonton; UCLA, Department of Zoology, University of California, Los 
Angeles; WAM, Western Australian Museum, Perth. 


Creedia alleni, new species 
Fig. | 


Holotype.—WAM P25808-005, 40.0 mm SL, probably a female, off Garden 
Island, near Fremantle, Western Australia, 32°15.7’S, 115°39’E, collected with 
beam trawl in 20 m, 29 June 1977. Bottom: sand and weed. 


30 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


eo 
5mm 


Fig. |. Creedia alleni, holotype, WAM P25808-005, 40 mm SL. A, Dorsal view of head region; 
B, Lateral view. 


Paratypes.—WAM P25346-028, 34.5 mm SL, off Gage Roads, Carnac Island, 
between Rottnest Island and Fremantle, Western Australia, 32°02’S, 115°40’E, 
collected with dredge, 30 June 1975, subsequently stained; WAM P25347-001, 
28.2 mm SL, same locality, 24 July 1975. 

Diagnosis.—A creediid with a I,3 pelvic fin (Fig. 2). All other creediids have 
the pelvic fins absent or I,4-5 (most I,5). Differs further from C. haswelli in 
having, on average, slightly fewer dorsal-fin rays (12 or 13 vs. 13-16), anal-fin rays 
(24 vs. 24 or more and usually 25 or 26), and vertebrae (41 or 42 vs. 42-45, usually 
43 or 44). The two tear-shaped epurals (observable in stained specimen only), 
may be unique in creediids (the epurals are relatively narrow in all other species 
with two). 

Description.—Morphometric and meristic data are given in Table |. Snout 
relatively elongate, sloping gradually in dorsal profile. Upper jaw with fleshy 
extension anterior to lower jaw; maxilla extending posteriorly to, or slightly be- 
yond vertical at center of eye; posterior tip of maxilla with well-developed notch 
(fork), lower lobe of fork longer than upper, upper lobe hidden under sheath 
when mouth closed. Bony dorsal projection at symphysis of lower jaw (Fig. 3). 
Lower jaw bordered by one row of pointed cirri of relatively uniform length (about 
17 per side in the holotype, 16 in larger paratype, and 11 in smaller paratype). 
Eyes dorsal; interorbital distance small (fleshy width about 5%c SL and bony width 
about half that); small, fleshy sheath covers lowermost part of eye. Tongue long 
and slender with slightly expanded, blunt, anterior tip. Ventral opercular flap 
extends far forward, overlapping branchiostegal membrane; posterior portion of 
gill cover overlaps base of pectoral fin. Branchiostegal rays seven (only six def- 
initely seen in holotype). Bone of gill cover splintered (clearly visible in unstained 


VOLUME 96, NUMBER 1 31 


1mm 1mm 


Fig. 2. Pelvic fin region of Creedia. A, Creedia alleni, paratype, WAM P25346-028, 34.5 mm SL, 
stained (right pelvic spine broken); B, C. haswelli, AMS IB.1114, 44 mm SL, New South Wales, 
cleared and stained. 


material with transmitted light), subopercle and interopercle heavily indented but 
preopercle bone very lightly incised and no splintering seen on opercle of stained 
paratype (Fig. 3). Body completely scaled and cheeks with at least some scales 
(most or all scales lost but scale pockets are present). Lateral line (clearly seen 
only in smaller paratype) arising at upper edge of gill cover, descending steeply 
at posterior portion of pectoral fin, and running parallel to and near ventral profile; 
tenth pored scale (at bottom of descending portion near tip of pectoral fin) and 
following scales on lateral line with elongate posterior lobes. Base of pelvic fins 
slightly anterior to base of pectoral fins; inter-pelvic fin distance very small (dis- 
tance between innermost rays about or less than 3%c SL). Dorsal, anal, pectoral, 
and pelvic rays unbranched. Caudal fin with nine branched rays (11 principal 
rays). Stained paratype with narrow neural and haemal spines in caudal peduncle 
region (similar to Limnichthys polyactis [Nelson 1979, fig. 3A] except that last 
haemal spine is also narrow); two large, contiguous tear-shaped epurals with apex 
at distal end, expanding proximally to broadly rounded proximal end; anterior 
epural broader, sloping strongly anteriorly at proximal end, distal third narrow, 
and anterior surface concave; posterior epural with shorter narrow distal part, 
sides gradually diverging proximally (epural shape clear only in the stained spec- 


32 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 3. Creedia alleni, paratype, WAM P25346-028, 34.5 mm SL, stained, right lower jaw and 
opercular region. Abbreviations: ANG, angular; ART, articular; DEN, dentary; IOP, interopercle; 
OP, opercle; POP, preopercle; SOP, subopercle. Gill cover membrane pulled down on ventral surface. 


imen and its radiograph). Stained paratype with single row of well-developed 
teeth along upper and lower jaws, ending well before anterior tip, and patch of 
well-developed teeth in middle of expanded anterior tip of premaxilla (unstained 
specimens appear similar); weakly-developed teeth appear to be present on vo- 
mer. 

No color pattern is apparent. 

Etymology.—The species is named after Gerald R. Allen, Curator of Fishes at 
the Western Australian Museum, who brought the specimens of the new species 
to my attention, in recognition of his many contributions to ichthyology. 

Discussion.—Creedia alleni is very similar to C. haswelli, and aside from the 
number of pelvic-fin rays, differs from other creediids in having the lowest number 
of dorsal-fin rays, and, possibly, in the shape of the epurals. 


Table 1.—Morphometric and meristic data of the three type specimens of Creedia alleni (WAM) 
and nine type specimens of Creedia partimsquamigera (first seven AMS and last two NMYV). Pro- 
portional measurements expressed as thousandths of standard length. * denotes decapitated specimen. 


C. alleni C. partimsquamigera 
Holo- Paratypes Holo- Paratypes 
type — type Seo SSS 

Standard length (mm) 40 35 28 55 67 59 57 55* 52 39 64 49 
Sex QY — — Q Q Q g 2 OY 6? 2 3? 
Body depth 72 70 78 61 63 61 60 ©661 — 5 SH 53 
Depth of caudal peduncle 34 35 41 35 32) 820 3038283 32 = 31 33 
Predorsal length 630 640 624 580 606 596 592 596 — 580 590 585 
Preanal length 446 459 450 460 456 462 451 438 — 448 466 462 
Pectoral fin length — 101 109 55.61. 59 S54. 54, il 9s Secale, 
Pelvic fin length 48 5] 58 35 39° 36 34 36 «#75 62> 33 69 
Head length 206 222 223 174 172 171 4170 173:«=163 «©1174 175 «169 
Head width 62 61 60 56" S9M2 37 55:9 53 SAS Ses 
Snout length 44 49 53 39 36 36 32 34 34 #38 «+36 34 
Length of orbit 31 35 32 22 21 22,20), <2 a t=, 1 22 18 20 
Dorsal-fin rays 13 12 12 16 16 = 15 15 Ib ils IG IGS) 
Anal-fin rays 24 24 24 25 Se DP 2 DS 2 2 25 
Pectoral-fin rays 12 12 12 13 | Zecepeieel) eed Aaah Bly abekalt2 ek eae « II) 12 
Pelvic-fin rays 1,3 I,3 1,3 I4 14 14 I1,4 1,4 14 1,4 #1,4 ~ 1,4 
Lateral-line scales c. 42 c. 42 c. 40 46 46 47 44 45 — 46 45 45 


Vertebrae 4] 41 42 45, 45)..46) 46) oe VAT ASG 


VOLUME 96, NUMBER 1 33 


Fig. 4. Creedia partimsquamigera, holotype, AMS 1.21420-001, 54 mm SL. A, Lateral view of 
head region; B, Lateral view. 


Creedia partimsquamigera, new species 
Fig. 4 


Holotype.—AMS 1.21420-001, 54.5 mm SL, ripe female, Coogee Beach, Syd- 
ney, New South Wales, 33°56’S, 151°16’E, 26 January 1980. Depth 10-15 m, in 
clean sand. 

Paratypes.—AMS 1.21420-005, 4 specimens, 52-67 mm SL (two specimens 
cleared and stained, including the smallest which was decapitated at capture), 
taken with the holotype. AMS I.22868-001, 2 specimens, 39 and 57 mm, Clovelly, 
Sydney, New South Wales, 33°53’S, 151°15’'E, 9 September 1981. NMV-A2229, 
2 specimens, 49 and 64 mm SL, Bondi Bay, Sydney, New South Wales, 33°54’S, 
151°17’E, 6 August 1980. All 9 specimens were collected by R. H. Kuiter with a 
handnet. 

Diagnosis.—A creediid with scales absent from anterior half of body except 
for the lateral line and a paired predorsal row, and 14—16 dorsal-fin rays. Differs 
further from C. haswelli and C. alleni in having a snout with a convex profile, 
the maxilla extending well behind the eye, the posterior tip of maxilla blunt, not 
forked, and the three or so elongate-most pectoral rays of males branched. 

Description.—Morphometric and meristic data are given in Table 1. Snout 
relatively short, strongly convex in dorsal profile. Upper jaw with fleshy extension 
anterior to lower jaw; maxilla extending posteriorly to well behind eye; posterior 
tip of maxilla blunt, notch weak if present, upper portion of maxilla hidden under 
sheath when mouth closed. Bony dorsal projection at symphysis of lower jaw. 
Lower jaw bordered by one row of blunt cirri, some alternating in length, about 
12-20 per side. Eyes dorsal; total interorbital width about 9-12%o SL, bony width 
about half that; fleshy sheath covers lowermost part of eye. Tongue long and 
slender. Ventral opercular flap extends far forward, overlapping branchiostegal 
membrane; posterior portion of gill cover overlaps base of pectoral fin. Bran- 
chiostegal rays 7. Bone of gill cover splintered, subopercle and interopercle heavi- 
ly indented, preopercle lightly indented, opercle lacks splintering. Scales present 
along lateral line, along base of dorsal fin and extending forward along midline 
in two adjacent rows to as far as about half distance from origin of dorsal fin to 
nape, and on body behind anterior portion of dorsal fin (usually about level of 
fifth or sixth ray); scales absent from anterior part of body except as noted. 
Lateral line arising at upper edge of gill cover, descending steeply near posterior 


34 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


tip of pectoral fin (behind in females and before in males), and running parallel 
to and near ventral profile; ventral lateral-line scales with elongate posterior lobes. 
Base of pelvic fins slightly anterior to base of pectoral fins; inter-pelvic fin distance 
very small, less than base length of fin. Dorsal, anal, pectoral, and pelvic rays 
unbranched except three or so elongate-most pectoral rays and perhaps elongate- 
most pelvic ray in males branched. Caudal fin usually with 9 branched rays. Single 
row of well-developed teeth along upper and lower jaws, ending well before 
anterior tip, and patch of well-developed teeth on relatively small expansion at 
anterior tip of premaxilla; vomerine teeth well-developed. 

Little to no color pattern is apparent in the preserved material. However, in 
color transparencies provided by R. H. Kuiter of the 64 mm paratype there are 
two stripes extending along the length of the body, one dorso-lateral and the 
other mid-lateral, and about 5 short, closely-set saddles just behind the nape. 

Etymology.—tThe specific name partimsquamigera is Latin (masculine) for partly 
scaled. 

Discussion.—Creedia partimsquamigera differs from the other species of Cree- 
dia in several characters (see Diagnosis). The absence of scales on much of the 
body is a feature similar to that found in the two nominal species of Chalixodytes 
Schultz. However, Creedia partimsquamigera has an abruptly descending lateral 
line unlike that of Chalixodytes (the key in Nelson 1978, is in error on this point) 
and is further similar to Creedia haswelli in the reduced number of dorsal-fin rays 
and having marked sexual dimorphism in pectoral- and pelvic-fin length (see later 
for evidence of this in C. haswelli). 

The type material of C. haswelli (and of C. clathrisquamis Ogilby, regarded as 
conspecific with C. haswelli) is from the Sydney area as is that of C. partim- 
squamigera. Denise S. Rennis has kindly confirmed for me that the four type 
specimens of C. haswelli and the one of C. clathrisquamis have completely scaled 
bodies. 


Other Australian creediids 


Creedia haswelli (Ramsay).—This species is known from southeastern Austra- 
lia from the Furneaux Islands off northeastern Tasmania (Scott 1969), northeast- 
ern Tasmania (Scott 1982), Victoria, New South Wales, and, on the basis of 
material not previously studied, from the vicinity of King Island in Bass Strait, 
and from western South Australia. 

Marked sexual dimorphism exists in the pectoral- and pelvic-fin length as de- 
termined from the large series from Bass Strait (NMV 2219-2228, 111 specimens). 
Nine specimens of each sex have the following fin lengths (in %c SL): females 
(3.5-4.9 cm SL), pectoral 90-102 and pelvic 45-54; males (3.0-4.9 cm SL), pec- 
toral 167-201 and pelvic 97-116. In the males the longest pectoral ray is the third 
or fourth dorsalmost while the longest pelvic ray is the outermost soft ray. Female 
C. haswelli thus have a pectoral fin length similar to that of Tewara cranwellae 
Griffin while in males it is more similar to that of Limnichthys fasciatus Waite, 
L. rendahli Parrott, and L. polyactis Nelson. Other than in C. partimsquamigera, 
such a marked sexual dimorphism in fin length is unknown in creediids. Scott 
(1969) noted serrations on the posterior lobe of the lateral-line scales. This feature 
shows some sexual dimorphism. In the NMV material (60 specimens examined) 


VOLUME 96, NUMBER 1 35 


most females have a smooth margin to the posterior lobe while most males have 
a serrated border on at least the lower margin (the serrations consist of small, 
isolated, flaplike projections which can be curled up and easily missed although 
some males do seem to lack them). A few females have a weakly serrated lower 
margin, similar to some males. Three hardened specimens from Tasmania (QVM 
1980/5/38, males as judged from the length of the paired fins) have a serrated 
ventral margin, similar to the NMV males. One male from South Australia (AMS 
1.10412, see below) has exceptionally long projections and at least one male from 
New South Wales (AMS 1I.18547-001) has moderately long projections on both 
the upper and lower margins. A few notches are also present on the anterior 
portion of the lobe in many specimens, both males and females, but occurrence 
of these notches was not studied in detail. 

Of 30 specimens radiographed, 6 have 42 vertebrae, 14 have 43, 9 have 44, and 
1 has 45. “ 

The South Australian material (AMS E1004, one specimen c. 36 mm, 32°31’S, 
133°18’E, south of St. Francis Island, 55 m, dorsal and anal fin rays not countable, 
and AMS I.10412, two specimens 37 and 38 mm, 32°35’S, 133°18’E, south of St. 
Francis Island, 64 m) has a pelvic fin of I,4, a dorsal fin with 14 rays, and an anal 
fin with 24 and 25 rays. 

Most specimens of Creedia haswelli, as with other creediids, are known from 
less than 100 m depth. One 40 mm ripe female (TFDA) collected off King Island, 
Bass Strait, however, was from 200 m. 

With the inclusion of C. alleni and C. partimsquamigera, the genus Creedia is 
diagnosed as follows: dorsal fin with 12—16 rays and anal fin with 24—28 rays; 
pelvic fin I,3 or 1,4; isolated ‘‘patch’’ of well-developed teeth on anterior tip of 
premaxillary bone; lateral line descending abruptly near tip of pectoral fin and 
running adjacent to anal fin base; lateral-line scales 40-47, those behind pectoral 
fin with posterior extension and not trilobate; marked sexual dimorphism in length 
of pectoral and pelvic fins (the condition in C. alleni is not known). 

Limnichthys fasciatus Waite.—Whitley (1945) erected the subspecies L. fas- 
ciatus major from one Western Australian specimen on the basis of it having only 
21 dorsal-fin rays and 24 anal-fin rays. In a previous study (Nelson 1978) I counted 
26 and 28 rays, respectively, in the same specimen but because it and three other 
specimens from Western Australia had relatively short predorsal distances I pro- 
visionally recognized the subspecies. An examination of a few characters in ad- 
ditional material of this species from Western Australia (AMNH 31323, three 
specimens 28-48 mm SL of five sent and nine in collection, Great Australian 
Bight, 34°56’S, 118°12'’E; UCLA W55-186, seven specimens, one of which is 
cleared and stained, 29-41 mm SL, Nancy Cove, Rottnest Island, 32°00’S, 
115°30’E, off Perth) did not reveal any marked differences in proportional mea- 
surements or meristic characters in comparison with material from eastern Aus- 
tralia and elsewhere although the predorsal distance is in the lower range for the 
species. The 10 specimens have the following characteristics: predorsal distance 
445475%o SL (& = 463); preanal distance 394-450 (« = 426); dorsal-fin rays 24— 
27 (& = 25.5); anal-fin rays 27-29 ( = 27.9); pectoral-fin rays 11-13 @ = 11.9); 
branched caudal-fin rays 8, lateral line descending gradually to end of base of 
anal fin; one to several rows of minute teeth on upper and lower jaws and ending 
well before tip and no teeth at tip of premaxilla (generally only one row for most 


36 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


of toothed portion in upper jaw); vomerine teeth numerous, minute, in two patches; 
about 15-21 cirri along margin of lower jaw (each side) with some cirri alternating 
from long to short along posterior part of jaw; some long cirri in three specimens 
of UCLA sample with two or three filaments off a wide tip, other UCLA and all 
AMNH specimens with pointed cirri (I have not examined the cirri in other L. 
fasciatus in detail but specimens from Lord Howe Island, UAMZ 3765, have 
pointed cirri); and cleared and stained specimen with two slender epurals and 
lacking preopercular splintering. The color pattern is present in the AMNH ma- 
terial and apparent, though faintly, in one UCLA specimen. The pattern is similar 
to that found in L. fasciatus elsewhere with bars extending ventrally from the 
dorsal saddles and approaching and usually joining expansions along the well- 
developed lateral band as shown in Nelson (1978, fig. 1C). Although there is no 
firm basis at present for recognizing a separate subspecies in Western Australia, 
further study is warranted of suspected differences between Western Australian 
material and that from elsewhere in the number of rows of teeth in the lower jaw 
and in the presence or absence of branching of the cirri bordering the lower jaw. 

Limnichthys donaldsoni Schultz.—Small specimens (8—19 mm SL) sent to me 
by D. F. Hoese from Yonge Reef, Lizard Island area, Queensland (15 specimens, 
AMS 1.19472-122, 14°35’S, 145°36’E), Great Detached Reef, Queensland (50 spec- 
imens, AMS I.20750-004, 11°39’S, 143°59’E), and Cape Melville, Queensland (21 
specimens, AMS I.20774-008, 14°10’S, 144°30’E) appear to represent L. donalad- 
soni. The few larger specimens from which accurate counts could be made have 
20 or 21 dorsal-fin rays and 24 or 25 anal-fin rays (relatively low compared to 
previous counts made on this species). The only specimens with clear markings 
have nine short dorsal saddles with no lateral band (unlike the pattern in L. 
fasciatus). This is the first evidence of L. donaldsoni in Australia and is a con- 
siderable southern extension in its known range. Limnichthys fasciatus is present 
at Lizard Island (AMS I.19473-189) and has a color pattern characteristic for that 
species (1.e., at least some dorsal saddles extending down to extensions on the 
well-developed lateral band; about 6—9 dorsal saddles in specimens 12-27 mm 
SL) and higher dorsal- and anal-fin ray counts than L. donaldsoni. 


Acknowledgments 


I thank G. R. Allen (WAM) and D. F. Hoese (AMS) for bringing specimens of 
Creedia in their care to my attention and making them available for description. 
I also thank M. F. Gomon (NMV), R. H. Green and E. O. G. Scott (QVM), P. 
Last (TFDA), D. S. Rennis (AMS), D. E. Rosen and C. Hutchings (AMNH), 
and J. Seigel and B. W. Walker (UCLA) for the loan of specimens. B. Hutchins 
(WAM) and D. F. Hoese read an earlier version of the manuscript and provided 
useful information. R. H. Kuiter kindly provided information on the Coogee 
Beach habitat and color transparencies of creediid material. I am grateful to K. 
A. Garbrah for his advice on Latin names. The figures were prepared, under 
supervision, by Diane Hollingdale. W. E. Roberts (UAMZ) provided museum 
assistance. The study was supported by grant No. A5457 of the Natural Sciences 
and Engineering Research Council of Canada. 


VOLUME 96, NUMBER 1 37 


Literature Cited 


Nelson, J. S. 1978. Limnichthys polyactis, a new species of blennioid fish from New Zealand, with 
notes on the taxonomy and distribution of other Creediidae (including Limnichthyidae).—New 
Zealand Journal of Zoology 5:351-364. 


—. 1979. Some osteological differences between the blennioid fishes Limnichthys polyactis and 


L. rendahli, with comments on other species of Creediidae.—New Zealand Journal of Zoology 
6:273-277. 

Scott, E. O. G. 1969. Observations on some Tasmanian fishes: Part XVI.—Australian Zoologist 
15(2): 160-177. 

——. 1982. Notes on fishes in the collection of the Queen Victoria Museum, Launceston. Pt. 
1.—Records of the Queen Victoria Museum, Launceston. No. 74, 31 pp. 


Whitley, G. P. 1945. New sharks and fishes from western Australia. Pt. 2.—Australian Zoologist 
11(1):1-42. 


Department of Zoology, The University of Alberta, Edmonton, Alberta T6G 
2E9, Canada. 


Note added in press.—One specimen of Creedia alleni (AMS 1.23416-001), with 
I, 3 pelvics, from 34°32'S, 115°01’E, and 5 specimens of what is probably C. has- 
welli (AMS 1.23412-001 to I.23415-001), with I, 4 pelvics counted in the 4 un- 
damaged ones, from 32°42’S, 131°27’E to 34°32’S, 121°16’E, were examined 


after this paper was in press. The two species are thus now known to be rela- 
tively close to one another. 


PROC. BIOL. SOC. WASH. 
96(1), 1983, pp. 38-49 


TELEOST FISH REMAINS (OSTEOGLOSSIDAE, 
BLOCHIIDAE, SCOMBRIDAE, TRIODONTIDAE, 
DIODONTIDAE) FROM THE LOWER EOCENE 
NANJEMOY FORMATION OF MARYLAND 


Robert E. Weems and Stephen R. Horman 


Abstract.—Six taxa of teleost fishes have been recognized from the Nanjemoy 
Formation of Maryland. So far, all collected remains have been fragmentary. The 
recognized taxa are: Brychaetus muelleri, Cylindracanthus rectus, Sarda del- 
heidi, Cybium sp., Triodon antiquus, and Kyrtogymnodon sp. No previous rec- 
ords of teleosts from this formation are known. 


Except for a preliminary report on the remains described herein (Horman and 
Weems 1976), no specific reports of teleosts from the Nanjemoy Formation of 
Maryland or Virginia are known, though Fallah (1964) referred to a Cylindracan- 
thus specimen collected from the Pamunkey Group, which could have come from 
either the Aquia or Nanjemoy Formation. Collecting during the last 15 years has 
yielded sparse but reasonably diagnostic fragmentary remains, which represent 
what could have been an abundant and diverse teleost fauna. Two or more bones 
from the same individual, fish or tetrapod, have never been reported from the 
Nanjemoy. This lack of even semi-articulated remains is in marked contrast to 
conditions in the underlying Paleocene Aquia Formation and the overlying Mio- 
cene Calvert Formation. The sparsity of identifiable fish bones in the Nanjemoy 
may be the result of a very low rate of clastic influx and a hard substrate, so that 
skeletons that fell to the sea bottom became totally disarticulated and broken up. 
Bryozoan colonies on some bones suggest that the remains lay on the sea bed 
for some time before burial. 

All teleost remains described herein were collected from the bluffs upriver and 
downriver from Popes Creek, Maryland (Fig. 1). In these bluffs, Clark and Martin 
(1901) recognized two members within the Nanjemoy Formation, a lower Pota- 
paco Member and an upper Woodstock Member (Fig. 2). Though separated in 
these bluffs by a prominent layer of calcareous nodules, the members are largely 
defined on faunal grounds, and to our knowledge, an ability to map these members 
outside of the vicinity of the Popes Creek bluffs on physical stratigraphic grounds 
has not been demonstrated. Because the base of the Nanjemoy (the contact with 
the underlying Marlboro Clay) and the top of the Nanjemoy (the unconformable 
contact with the overlying basal phosphate bed of the Calvert Formation) are 
mappable stratigraphic contacts, the Nanjemoy as a whole is a mappable strati- 
graphic unit (Glaser 1971; Teifke 1973). Our work was done in the type area 
where the marker horizon between the two Nanjemoy members is readily seen; 
hence their names are retained for this report. Elsewhere, the members might be 
only biostratigraphic subdivisions of the Nanjemoy. 

Only the uppermost 15 feet of the Potapaco Member is exposed above Popes 
Creek, so we have no basis for disputing Clark and Martin’s (1901) estimate that 


“VOLUME 96, NUMBER 1 39 


Fig. 1. Location of the bluffs along the Potomac River near Popes Creek, Maryland (ruled area). 


this member is about 70 feet thick. Our own estimates on the thickness of the 
Woodstock Member, however, suggest that this member as Clark and Martin 
defined it at Popes Creek is 80 feet thick in the outcropping area instead of the 
50 to 60 feet they reported (Clark and Martin 1901:66). Curiously, their diagram- 
matic column of the Pamunkey Group indicates a thickness for the Woodstock 
of about 80 feet, suggesting that the published thickness may have been a typo- 
graphical error. According to our estimates, Popes Creek marks the spot at which 
50 feet of the Woodstock has passed below sea level. North of Popes Creek, a 
thin and apparently laterally impersistent layer of coarse sand and phosphate 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


40 


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Fig. 3. Known range of some species of vertebrates collected from the Nanjemoy Formation at 
Popes Creek, Maryland. Except for Myliobatis, Aetobatis, Ginglymostoma, and Odontaspis, which 
are relatively abundant, dots represent single collected specimens. 


—_— 


Fig. 2. Stratigraphic column of the Pamunkey Group (from Clark and Martin 1901, and Gibson 
et al. 1980), showing the age ranges of the various formations and groups. MYA = millions of years 
ago. 


42 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


pebbles offers the only well defined datum from which thickness estimates can 
be based. This bed is considered to be 20 feet above the base of the Woodstock 
Member as defined by Clark and Martin (1901). South of Popes Creek, the only 
persistent marker bed is a thin, 1-4 inch thick layer of carbonaceous plastic light- 
gray clay 76 feet above the base of the Woodstock and 4 feet from its top as 
exposed. Specimens of the large nautiloid, Hercoglossa tuomeyi Clark and Mar- 
tin, are known only from an interval 55—70 feet above the base of the Woodstock. 
Figure 3 portrays graphically the estimated position of these horizons, as well as 
the position of the teleost remains described below and some other vertebrate 
remains. 

Besides teleosts, rare fragmentary remains of crocodiles, turtles, and the “‘sea 
snake’ Paleophis are found in the Nanjemoy. Teeth of sharks and skates are 
more common than other vertebrate remains. Teeth of the shark Otodus obliquus 
Agassiz occur throughout the Aquia Formation and Potapaco Member, and those 
of the shark Procarcharodon auriculatus (de Blainville) are found throughout the 
Woodstock Member, but teeth of the two species do not occur together. The 
serrated tooth margin of P. auriculatus is the only difference between the two 
species; hence, these forms may represent a single lineage. If so, the two forms 
are good index fossils for distinguishing each respective member. In addition, the 
ray Aetobatis irregularis Leriche has not been reported in the Potapaco Member 
even though it is common throughout the Woodstock Member. Vertebrae of 
Paleophis are rare; only P. virginianus Lynn is present in the Potapaco, whereas 
the Woodstock vertebrae are referrable to P. grandis (Marsh). Thus, there seems 
to be a detectable vertebrate biostratigraphic break between the Potapaco and 
the Woodstock comparable to that found in the invertebrate assemblages (Clark 
and Martin 1901), though the phosphate pebble bed 20 feet above the basal con- 
cretion bed seems a more reasonable location for a time break. This break prob- 
ably affects the teleost distributions as well, but the teleost sample is far too 
sparse for any prediction to be made on which species might be confined to one 
member or the other. 

On the basis of the first appearance of Procarcharodon auriculatus, Leriche 
(1943) wanted to correlate the Woodstock with the Lutetian (Middle Eocene) of 
Europe, but since then P. auriculatus has been reported from the Ypresian (Low- 
er Eocene) of Britain as far down as approximately the base of nannoplankton 
zone NP13 (Hooker ef al. 1980). On the basis of microfossil correlations, the 
Woodstock recently has been placed in the slightly older lower to middle Ypresian 
fossil zones NP11 and 12 (Gibson et al. 1980). Since control on the boundaries 
of Paleogene nannoplankton zones in Britain is not good (Hooker et al. 1980), 
this slight difference in the time of first appearance for P. auriculatus on either 
side of the Atlantic may be more apparent than real. If it is real, then P. auric- 
ulatus appeared slightly earlier in the western Atlantic than in the eastern Atlan- 
tic. In either case, an Ypresian age for the Woodstock is compatable with all 
available data. The underlying Marlboro Clay of late Paleocene and early Eocene 
age (Gibson ef al. 1980) overlies the Aquia Formation, which has been well 
documented as Late Paleocene in age (Loeblich and Tappan 1957; Bybell and 
Govoni 1977; Gibson et al. 1980). Thus, these data indicate that the Potapaco 
Member also must be Early Eocene (Ypresian) in age. 


VOLUME 96, NUMBER 1 43 


Fig. 4. Right dentary of Brychaetus muelleri in external (A), internal (B), and dorsal (C) aspects. 
Length 17 cm. 


Order Osteoglossiformes 
Family Osteoglossidae 
Brychaetus muelleri Woodward 


Material.—Right dentary containing 17 teeth and 8 empty alveoli (USNM 
265383), Fig. 4. 
Locality.—Bluff 100 yards south of Popes Creek, Maryland, within uppermost 


44 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 5. A. Rostral fragment of Cylindracanthus rectus. Length 5.25. cm. B—F. Dentary of Triodon 
antiquus in anterior (B), posterior (C), right-lateral (D), ventral (aboral) (E), and dorsal (oral) (F) 
aspects. Length along midline, 6 mm; width at rear of jaw rami, 10 mm. 


10 feet of the Woodstock Member, Nanjemoy Formation (‘‘Zone 17’’ of Clark 
and Martin 1901), coll. John Glaser, 1972. 

- Discussion.—This large dentary with its hollow, laterally wide, anteroposte- 
riorly compressed, and lingually recurved teeth is highly distinctive and readily 
referrable to B. muelleri (compare with Casier 1966). This teleost is by far the 
largest known from the Nanjemoy and probably was one of the most formidable 
marine carnivores of its time. Phareodus, a related osteoglossid from the Eocene 
Green River shales, is only about one-sixth as large, has a proportionally deeper 


VOLUME 96, NUMBER | 45 


jaw, and teeth that are not nearly so lingually recurved. Brychaetus has not been 
previously reported from the Lower Eocene of North America (Horman and 
Weems 1976). 
Order Perciformes 
Family Blochiidae 
Cylindracanthus rectus (Agassiz) 


Material.—Two rostral fragments, one of which (USNM 265384) is shown in 
Fig. 5(A). 

Locality.—(1) Bluff % mile north of Popes Creek, Maryland, 4 feet above 
beach, Woodstock Member, Nanjemoy Formation, coll. Robert E. Weems, 1973; 
(2) USNM 265384, bluff ’2 mile south of Popes Creek, Maryland, 2 feet above 
beach, Woodstock Member, Nanjemoy Formation, coll. Robert E. Weems, 1975. 

Discussion.—Fragmentary rostra referred to this genus are fairly common in 
lower Tertiary sediments of North America and Europe (for example, Fallah 
1964). Several species have been named, but most are based on such fragmentary 
material that their validity is questionable. The specimens described here are 
fully comparable with C. rectus, the first named species of the genus, and are 
thus placed in that taxon. Until a great deal more well-preserved material becomes 
available, it is impossible to tell whether this is a discrete taxon or only a catch- 
all category for rostral fragments of any of several closely related fishes. 


Family Scombridae 
Sarda delheidi (Leriche) 


Material.—Right dentary (USNM 265383) with 6 teeth and at least 10 empty 
alveoli (Fig. 6A—C). 

Locality.—Bluff 1 mile south of Popes Creek, Maryland, | foot above beach 
and 15 feet below gray carbonaceous clay bed, Woodstock Member, Nanjemoy 
Formation (‘Zone 17’ of Clark and Martin 1901), coll. Robert E. Weems, De- 
cember 1973. 


Cybium sp. 


Material.—Three isolated teeth (Fig. 6D-F). 

Localities.—1) USNM 265389, bluff 2 miles north of Popes Creek, Maryland, 
coll. Robert E. Weems, 1973. 2) USNM 265385, USNM 265386, bluff 1 mile north 
of Popes Creek, Maryland, | foot above beach in phosphate lag deposit, Wood- 
stock Member, Nanjemoy Formation (“‘Zone 16”’ of Clark and Martin 1901), coll. 
Stephen R. Horman and Robert E. Weems, July 1968. 

Discussion.—Isolated teeth of this sort have been traditionally assigned to Cy- 
bium, (compare with Casier 1966, Leriche 1905), and occasionally even species 
level identifications have been attempted. The teeth illustrated here are not cer- 
tainly identifiable to species but are fully comparable with teeth generally assigned 
to this genus. 


Order Tetraodontiformes 
Family Triodontidae 
Triodon antiquus Leriche 


Material.—Fused dentary beak (USNM 265387) with dental battery (Fig. 
SB-F). 


46 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 6. A-—C. Right dentary of Sarda delheidi in dorsal (oral) (A), external (B), and internal (C), 
aspects. Length 9.4 cm. D-F. Teeth referrable to Cybium sp. Height about 8 mm. G—H. Dental plate 
of Ayrtogymnodon sp. in internal (G) and external (H) aspects. Length 9 mm, width 15 mm. 


Locality.—Bluff 1 mile north of Popes Creek, Maryland, 3 feet above beach 
and 2 feet above phosphate lag deposit, Woodstock Member, Nanjemoy For- 
mation (“‘Zone 16” of Clark and Martin 1901), coll. Robert E. Weems and Stephen 
R. Horman, July 1968. 

Discussion.—This fused dentary beak obviously belongs among the tetraodon- 
tiform fishes. The Molidae and Diodontidae, however, lack the scalelike teeth 


VOLUME 96, NUMBER 1 47 


covering the beak margin, and the diodontidae have the tooth plates stacked 
imbricately, not arranged anteroposteriorly as in the Triodontidae. The only living 
species of this family is Triodon bursarius (Tyler, 1962). Fossil species of this 
genus are Triodon antiquus Leriche (1905) and 7. cabindensis Leriche (1920). 
Our specimen of Triodon has a beak nearly as long as wide, differing markedly 
from the extremely laterally elongate beak of T. cabindensis. In general propor- 
tions it is like 7. antiquus, being about the same size and shape. Most specimens 
of T. antiquus have beaks notably wider than long, however, whereas in this 
specimen the proportions are nearly equal, thus giving an exceptionally sharp 
and angular appearance to this beak when seen from dorsal aspect. This appears 
to have resulted only from proportional changes related to growth. Most figured 
specimens of 7. antiquus (Leriche 1905; Casier 1946, 1960) are smaller than this 
specimen. When specimens are compared by absolute size, instead of relative 
size, the beak curvatures are quite comparable, suggesting that the differences 
in apparent curvature may be due to the greater individual age of our specimen 
than that of most of the figured specimens from Europe, whose jaw rami had not 
grown to be so elongate. Only two of the European specimens reach a size 
comparable to our specimen. In most of the figured specimens of T. antiquus, 
only juveniles have a single pair of teeth on the inferior oral surface. By the time 
specimens reach sizes approaching that of our specimen, one or more pairs of 
lateral teeth erupt as well. Thus our specimen is unusual in this respect, but in 
view of the highly variable nature of the dentition in figured specimens of T. 
antiquus, it is not truly distinctive when based on only a single specimen. There- 
fore, unless enough material becomes available to allow a statistical comparison 
between the European and American populations, this specimen is best referred 
to the highly variable Triodon antiquus. 

This is the first record of this family from either North America or the Western 
Hemisphere. Today, the family is restricted to the Indian Ocean and in the past 
is known to have ranged into Europe and West Africa only during the Eocene. 
Apparently, at that time the family also managed to cross the North Atlantic into 
North America. If McKenna (1972) is correct in postulating a landbridge between 
Europe and North America through Early Eocene time, this would have afforded 
a warm Shallow dispersal route for this family, whose distribution today suggests 
that it may be restricted to warm waters. 


Family Diodontidae 
Kyrtogymnodon sp. 


Material.—Oral tooth plate, USNM 265388, (Fig. 6G—H, containing a pair of 
tooth batteries each containing stacks of three teeth. 

Locality.—2 miles north of Popes Creek, Maryland, along a low bluff at the 
mouth of Nanjemoy Creek in the uppermost 10 feet of the Potapaco Member of 
the Nanjemoy Formation (“‘Zone 15”’ of Clark and Martin 1901), coll. Robert E. 
Weems, 1973. 

Discussion.—TYavani (1955) reviewed the known diodontids and arranged them 
within four genera. Although his suggestions concerning the definitions of genera 
are accepted here, it is by no means certain that species characters can be rec- 
ognized from a single pair of fused plates. Moreover, it may well be that the total 
number of presently described species exceeds the true numbers of species rep- 


48 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


resented, since individual variants may have been separately described. There- 
fore, the pair of fused dental plates illustrated here are merely designated Kyr- 
togymnodon sp. until better remains permit a specific diagnosis. This plate is 
assigned to Kyrtogymnodon because wear is restricted almost wholly to the cen- 
tral region of the most-erupted tooth. This suggests that the entire tooth battery 
was stacked nearly vertically so that only one tooth was worn at a time. In 
Progymnodon, Oligodiodon, and Diodon the batteries are stacked obliquely so 
that the edges of at least several teeth are functioning at the same time. If this 
assignment to Kyrtogymnodon is correct, this is by far the oldest known occur- 
rence of this genus. Previously it has only been described from the Pliocene 
(Tavani 1955). 


Summary 


The known teleost remains from the Nanjemoy Formation can be stratigraph- 
ically arranged as follows: 


Woodstock Member—Cylindracanthus rectus (Agassiz) 
Triodon antiquus Leriche 
Brychaetus muelleri Woodward 
Sarda delheidi (Leriche) 
Cybium sp. 

Potapaco Member— Kyrtogymnodon sp. 
Cybium sp. 


These species probably represent only a small fraction of the total number of 
teleost species originally present; conditions during deposition seem to have de- 
stroyed all remains except those from species that had exceptionally durable 
skeletal elements. All remains are fragmentary, and no associated skeletal ma- 
terials are known. 


Literature Cited 


Bybell, L. M., and D. C. Govoni. 1977. Preliminary calcareous nannofossil zonation of the Bright- 
seat and Aquia Formations (Paleocene) of Maryland and Virginia—stratigraphic implications.— 
American Association of Petroleum Geologists, National Meeting (Washington, D.C.), Program 
and Abstracts, p. 67. 

Casier, E. 1946. La faune ichthyologique de 1’ Ypresien de la Belgique.—Memoires du Musée Royal 
d Histoire Naturelle de Belgique 104:1—267. 

—. 1960. Note sur la collection des poissons Paléocenes et Eocenes de l’Enclave de Cavinda 
(Congo).—Annales du Musée Royal du Congo Belge (3) 1 (2):1—47. 

——. 1966. Faune ichthyologique du London Clay.—British Museum (Natural History): London. 
2 vols., 496 pp. 

Clark, W. B., and G. C. Martin. 1901. The Eocene deposits of Maryland.—Maryland Geological 
Survey, Eocene, pp. 19-92. 

Fallah, W. 1964. Cylindracanthus from the Eocene of the Carolinas.—Journal of Paleontology 38: 
128-129. 

Gibson, T. G., G. W. Andrews, L. M. Bybell, N. O. Frederiksen, T. Hansen, J. E. Hazel, D. M. 
McLean, R. J. Witmer, and D. S. van Nieuwenhaise. 1980. Geology of the Oak Grove Core, 
Part 2: Biostratigraphy of the Tertiary strata of the core.—Virginia Division of Mineral Re- 
sources, Publication 20:14—30. 

Glaser, J. D. 1971. Geology and mineral resources of southern Maryland.—Maryland Geological 
Survey, Report of Investigations 15:1-84. 


VOLUME 96, NUMBER 1 49 


Hooker, J. J., A. N. Insole, R. T. J. Moody, C. A: Walker, and D. J. Ward. 1980. The distribution 
of cartilaginous fish, turtles, birds and mammals in the British Paleogene.—Tertiary Research 
3(1): 1-12. 

Horman, S. R., and R. E. Weems. 1976. Eocene fish and reptiles from the Nanjemoy Formation 
in Maryland.—Geological Society of America, Abstracts of 25th Annual Northeastern/South- 
eastern Section 8(2):198. 

Leriche, M. 1905. Les poissons Eocene de la Belgique.—Extrait Memoire Musée Royal d’ Histoire 
Naturelle Belgique 3:49-228. 

———. 1920. Notes sur la Paleontologie du Congo. III. Note préliminaire sur des poissons nouveaux 
du Paléocene et de l Eocene de la region cotiére du Congo.—Revue Zoologique Afrique 8:85. 

———. 1943. Le synchronisme des formations Tertiares des deux cotes de |’Atlantique, d’apres 
leur faune ichthyologique.—Comptes Rendue de Séances de 1’ Academie des Sciences 210:589— 
592. 

Loeblich, A. R., and H. Tappan. 1957. Correlation of the Gulf and Atlantic Coastal Plain Paleocene 
and lower Eocene formations by means of planktonic Foraminifera.—Journal of Paleontology 
31:1109-1 137. 

McKenna, M. C. 1972. Was Europe connected directly to North America prior to the Middle 
Eocene?—In Dobzhansky, Hecht, and Steere eds. Evolutionary Biology, vol. 6, pp. 179-188. 
New York: Appleton-Century-Crofts. 

Tavani, G. 1955. Osservazioni su alcuni Plectognathi (Gymnodonti).—Atti Societe Toscana Scienze 
Naturalle Memorie ser. A, 62:177—200. 

Teifke, R. H.” 1973. Stratigraphic units of the Lower Cretaceous through Miocene series, in Geo- 
logical Studies, Coastal Plain of Virginia.—Virginia Division Mineral Resources Bulletin 83: 
1-78. 

Tyler, J. C. 1962. Triodon bursarius, a plectognath fish connecting the scleroderms and gymno- 
donts.—Copeia 4:793-801. 


(REW) United States Geological Survey, Stop 928, Reston, Virginia 22092; 
(SRH) Naval Surface Weapons Laboratory, Dahlgren, Virginia 22448. 


PROC. BIOL. SOC. WASH. 
96(1), 1983, pp. 50-58 


REDESCRIPTION OF THE BIGEYE SHINER, 
NOTROPIS BOOPS (PISCES: CYPRINIDAE) 


Brooks M. Burr and Walter W. Dimmick 


Abstract.—The wide-ranging yet poorly known bigeye shiner, Notropis boops, 
is redescribed from examination of 567 specimens from throughout the range of 
the species. Significant geographic variation was not found in any of the 20 mor- 
phological characters examined. Numbers of lateral-line scales and caudal ver- 
tebrae were slightly higher in the northern and eastern parts of the species’ range. 
Populations of N. boops have been severely decimated in both Ohio and Illinois 
due to excessive siltation from poor agricultural practices; it is a common species 
throughout the central portion of its range. 


The bigeye shiner, Notropis boops Gilbert, is a mostly common inhabitant of 
the central Mississippi basin, although it is disappearing or rare in some areas on 
the edges of its range, e.g., Ohio (Trautman 1981), Illinois (Smith 1979), and 
Alabama (Ramsey 1976). Since its original description (Gilbert 1884), little infor- 
mation has been published on the systematics of N. boops, except for its inclusion 
in atlases, checklists, and state fish books. Recently, Gilbert (1978) listed the type 
material of Notropis boops, Smith and Hocutt (1981) reported on variation in 
pharyngeal tooth formulae in Missouri, and Lehtinen and Echelle (1979) analyzed 
the reproductive cycle of the species in Oklahoma. 

Because of its rather wide range on both sides of the Mississippi River and the 
paucity of systematic information available on the species, we undertook a study 
of geographical variation in N. boops. Examination of 20 morphological char- 
acters on 567 specimens from throughout the range of the species revealed that 
geographic variation is minimal. The objectives of this paper are to redescribe 
N. boops and to map and discuss its distribution. 


Methods 


Counts and measurements followed those described by Hubbs and Lagler (1964). 
In examination of the cephalic lateral line, the terminology, abbreviations, and 
counting procedures of Snelson (1971) were followed. Vertebral counts were 
made from radiographs. The first vertebra bearing a well-developed haemal spine 
was considered to be the first caudal element; all those anterior were considered 
trunk vertebrae. 7 

All meristic characters were analyzed initially for sexual or geographic variation 
in minor drainages. When no significant sexual, intra- or interdrainage variation 
was apparent, the data were pooled into two major groups of populations—those 
occurring east of the Mississippi River and those occurring west of the river. 


Material Examined 


The following material was examined. Numbers of specimens counted or mea- 
sured are in parentheses. Institutional abbreviations are identified in Acknowl- 


VOLUME 96, NUMBER | 51 


edgments. Complete collection data are on deposit at the Department of Zoology, 
Southern Illinois University at Carbondale. 


Notropis boops Gilbert 


OHIO RIVER BASIN. Scioto R. drainage.—OH: Pike Co.: OSU 12334 (10). 
Scioto Co.: UMMZ 86011 (2). Kentucky R. drainage.—KY: Franklin Co.: UMMZ 
144395 (3). Lincoln Co.: INHS 79062 (10). Kinniconick Cr.—KY: Lewis Co.: UL 
5444 (5). Harrods Cr.—KY: Oldham Co.: UL 4779 (5). Salt R.-Rolling Fork 
drainage.—KY: Anderson Co.: UL 3175 (10). Jefferson Co.: UL 4774 (6). Mad- 
ison Co.: UMMZ 125060 (4). Shelby Co.: UL 5372 (6). Blue R.—IN: Washington 
Co.: OSU 28646 (9). Green R. drainage.—KY: Casey Co.: UMMZ 169436 (1). 
Muhlenberg Co.: SIUC 1915 (10). Wabash R. drainage.—IN: Carrol Co.: OSU 
27890 (2). Hamilton Co.: OSU 30104 (9). Lawrence Co.: UMMZ 167923 (3). 
Montgomery Co.: OSU 27169 (5). Parke Co.: UMMZ 144536 (1). Putnam Co.: 
OSU 27500 (8). Shelby Co.: OSU 29861 (10). IL: Edgar Co.: INHS 2937 (8). 
Shelby Co.: INHS 21720 (6). Vermilion Co.: INHS 12241 (8). Cumberland R. 
drainage. —IN: Smith-Wilson Co.: UT 44.249 (20). Williams Co.: INHS 83116 
(1); UMMZ 175215 (10). KY: Cumberland Co.: INHS 78328 (10). Tennessee R. 
drainage.—AL: Jackson Co.: AU 12003 (10). Limestone Co.: UMMZ 200821 (2); 
UMMZ 200870 (1). TN: Bedford Co.: UMMZ 121290 (10); UT 44.709 (21). Benton 
Co.: UT 44.964 (4). KY: Calloway Co.: SIUC 291 (4); SIUC 308 (10); SIUC 383 
(5); SIUC 388 (1). Marshall Co.: SIUC 1530 (6). 

MISSISSIPPI RIVER BASIN. Aaskaskia R. drainage.—IL: Moultrie Co.: 
INHS 8915 (10). Cuivre R. drainage.—MO: Pike Co.: UMMZ 149205 (10). Mer- 
amec R. drainage.—MO: Franklin Co.: SIUC uncat. (5). Gasconade Co.: UMMZ 
148339 (10). trib., Mississippi R.—MO: Perry Co.: UMMZ 149843 (8). Miller 
Cr.—IL: Alexander Co.: INHS 6047 (10). St. Francis R. drainage.—MO: Wayne 
Co.: SIUC uncat. (6). White R. drainage.—MO: Butler Co.: UT 44.942 (10). 
Carter Co.: UMMZ 193194 (5). Ripley Co.: UT 44.1826 (10). Shannon Co.: SIUC 
uncat. (8). Taney Co.: UMMZ 151181 (10). Clark Natl. Forest: UMMZ 117265 
(8). AR: Madison Co.: UT 44.897 (9). Seary Co.: UMMZ 123532 (6). Van Buren 
Co.: INHS 123570 (10); INHS 81072 (10). 

MISSOURI RIVER BASIN. Gasconade R. drainage. —MO: Gasconade Co.: 
INHS 80497 (3). 

ARKANSAS RIVER BASIN. Caney R. drainage.—KS: Chautauqua Co.: KU 
14424 (9). Grouse Cr.—Cowley Co.: KU 8616 (8). Neosho R. drainage.—KS: 
Cherokee Co.: KU 3192 (10); UMMZ 144939 (3); UMMZ 155100 (3); UMMZ 
155175 (1); UMMZ 160379 (3). OK: Delaware Co.: UMMZ 103170 (10). Illinois 
R. drainage.—AR: Washington Co.: INHS 82466 (10). OK: Adair Co.: UT 48.1848 
(3). Sequoyah Co.: UMMZ 137825 (6). trib., Arkansas R.—AR: Crawford Co.: 
UMMZ 123741 (10); UMMZ 123826 (10). Fourche La Fave R. drainage.—AR: 
Yell Co.: INHS 81092 (10). 

RED RIVER BASIN. Blue R. drainage.—OK: Johnston Co.: OUMZ 30478 
(10); OUMZ 40205 (4); UMMZ 156760 (4). Kiamichi R. drainage. —OK: LeFlore 
Co.: UMMZ 80947 (6). Little R. drainage. —OK: McCurtain Co.: INHS 80736 
(10). AR: Polk Co.: UMMZ 170884 (10). Saline R. drainage.—AR: Sevier Co.: 
UMMZ 128088 (3). Ouachita R. drainage.—AR: Polk Co.: INHS 81032 (10). 
Montgomery Co.: SIUC 2494 (5). Saline Co.: SIUC 3645 (2); SIUC 3700 (5). 


52 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 1. A, N. boops, 54 mm SL female (SIUC 2494), Caddo River, Montgomery County, Arkan- 
sas, 9 October 1981; B, N. xaenocephalus, 55 mm SL male (INHS 75095), Stamp Creek, Bartow 
County, Georgia, 9 June 1976. 


Notropis xaenocephalus (Jordan) 


MOBILE BAY BASIN. Etowah R. drainage.—GA: Bartow Co.: INHS 75095 
(10). Oostanaula R. drainage.—GA: Murray Co.: SIUC 2895 (10); SIUC 2884 
(2). GA: Gordon Co.: SIUC 2904 (5). 


Notropis boops Gilbert, 1884 
Fig. 1 


Types.—Syntypes of N. boops were collected by C. H. Gilbert from Salt Creek, 
Brown County, Indiana (10 specimens), and from Flat Rock Creek, Rush County, 
Indiana, by W. P. Shannon (about 30 specimens) (Gilbert 1884). C. R. Gilbert 
(1978) found and listed the following syntypes (numbers of specimens and their 
range in SL are in parentheses): Salt Creek—USNM 34982 (3, 44-46), MCZ 35961 
(1, 54); 6 specimens not located. Flat Rock Creek—CAS-SU 3794 (20, 47-61); 
about 10 specimens not located. Some doubt exists regarding the status of the 
CAS-SU specimens because nothing in the jar indicated they were types and 
Gilbert was listed as the collector. Gilbert (1978) recommended that a lectotype 
be chosen from among the Salt Creek specimens. A lectotype of N. boops (45.6 
mm SL) is herein selected and retains the original number USNM 34982. The 
lectotype conforms to the characters in the description that follows. The remain- 


VOLUME 96, NUMBER 1 53 


Table 1.—Proportional measurements (expressed in thousandths of SL) for Notropis boops in 
breeding condition summarizing sexual dimorphism. Specimens measured were 45-58 mm SL and 
were from the following drainages: Wabash (INHS 2937), Tennessee (UMMZ 200821), Mississippi 
(INHS 6047), White NHS 81072), Arkansas (INHS 81092), and Ouachita (INHS 81032, 82466). 


Character Sex n Range x ’ t-Value P 
Head length M 31 239-287 254 0.3536 N.S. 
F 19 245-266 253 
Head width M 3] 121-152 133 1.5504 N.S. 
F 19 124-140 130 
Head depth M 30 138-176 150 1.1960 N.S. 
F 19 138-156 148 
Eye diameter M 31 72-94 82 0.7226 N.S. 
F 19 76—96 83 
Snout length M 3] 58-71 65 1.8892 N.S. 
F 19 56-69 63 
Upper jaw length M 30 48-76 62 0.3077 N.S. 
F 19 53-72 61 
Bony interorbital length M 31 52-75 64 1.8051 N.S. 
F 19 53-72 62 
Anal fin length M 3] 139-191 167 2.7872 <.01 
F 19 139-182 159 
Dorsal fin length M 3] 138-267 228 0.7204 N.S. 
F 19 194-239 223 
Pelvic fin length M 31 149-184 166 4.3382 <.005 
F 19 143-175 155 
Pectoral fin length M 31 167-218 192 5.2201 <.005 
F 19 151-199 178 
Preanal length M 30 63 1-688 671 3.2045 <.005 
F 19 672-702 681 
Prepelvic length M 30 461-533 485 1.9223 N.S. 
F 19 462-510 492 
Predorsal length M 3] 469-553 49] 1.8442 N.S. 
F 19 472-523 499 
Postdorsal length M 31 500-609 532 1.5905 N.S. 
F 19 485-545 524 
Body width M 31 112-150 128 0.8994 N.S. 
F 19 113-150 131 
Body depth M 31 127-277 208 1.4403 N.S. 
F 19 130-256 223 
Caudal ped. depth M 3] 82-103 89 0.1100 N.S. 
F 19 79-94 87 
Caudal ped. length M 3] 113-256 220 0.3516 N.S. 
F 19 122-241 217 


ing syntypes now become paralectotypes bearing their original catalog numbers 
except those from USNM 34982 which now bear catalog number USNM 232410. 

Diagnosis.—A moderate-sized species of Notropis (largest specimen measured 
is 66 mm SL); peritoneum black; intestine short (76.5—98.0% SL) with a single 
lengthwise loop; eye large (less than 3.5 times into head); anal-fin rays usually 8; 
pharyngeal teeth usually 1,4—4,1; no breeding colors. 


54 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Table 2.—Counts of lateral-line scales in Notropis boops. 


Region 34 35 36 37 38 39 40 n x 
East of Mississippi R. 23 9] 86 17 3 ] 221 36.5 
West of Mississippi R. 3 25 125 81 18 225 35.9 


Description.—Proportional measurements are shown in Table |. General body 
shape and pigmentary features are illustrated in Fig. |. 

Dorsal-fin rays 7(1), 8(324). Anal-fin rays 7(5), 8(488), 9(24); « = 8.0. Left pec- 
toral-fin rays 13(5), 14(61), 15(270), 16(160), 17(17), 18(1); = 15.2. Pelvic-fin rays 
7-6(1), 7-7(6), 7-8(13), 8-8(474), 8-9(8), 9-9(5). Principal caudal-fin rays 18(9), 19(449), 
AV ZICGs ae = 120. 

Lateral-line scales 34—40 (Table 2). Body-circumference scales 21(7), 22(156), 
23(144), 24(99), 25(37), 26(31), 27(2); = 23.2. Body-circumference scales above 
lateral line 10(6), 11(360), 12(63), 13(47) ( = 11.3); below lateral line 8(4), 9(189), 
10(160), 11(119), 12(4) ( = 9.8). Predorsal scale rows 10(1), 11(3), 12100), 13(239), 
14(119), 15(36); « = 13.2. Caudal-peduncle scale rows above lateral line 5(502), 
6(6) (x = 5.0); below lateral line 5(506), 6(1), 711) @ = 5.0). Breast and nape 
completely scaled. Total gill rakers on first arch (all rudiments counted) 8(2), 9(6), 
OD), Wie ese = Do7/- 

Numbers of trunk, caudal, and total vertebrae are shown in Table 3. Pharyngeal 
teeth (sampled from throughout range) 1,4—4,0(6); !,4—4,1(42). Teeth with 
prominent terminal hooks, but narrow grinding surfaces. 

Lateral line on body complete to caudal base. Supratemporal canal broadly 
interrupted at dorsal midline; ST pore counts 2,2(20). Supraorbital canal without 
interruptions and not joining infraorbital canal behind eye; SO pore counts 7(5), 
8(12), 9(2), 10(1); x = 8.0. Infraorbital canal complete; IO pore counts 10(2), 11(5), 
12(12), 13(1); = 11.6. Preoperculomandibular canal without interruptions; POM 
pore counts 9(7), 10(11), 11(2); « = 9.8. 

Peritoneum black ventrally, silvery with scattered melanophores on dorsal sur- 
face of body cavity. Intestine short with a single lengthwise loop (Type I of 
Kafuku 1958). In 10 adults 48-59, x = 54 mm SL, ascending section 22.8—32.8, 
x = 28.6% SL; total intestinal length 76.5—98.0, x = 87.8% SL. Gas bladder two- 
chambered; total length 31.6—37.6, x = 33.8% SL, posterior chamber length 21.2— 
23.7, X = 22.7% SL. Peritoneal tunic covers anterior chamber. Pneumatic duct 
attached to dorsal surface of anterior chamber. Bladder lacks spiral markings or 
Striations. 

Distinct black lateral stripe about 1—1'4 scale rows wide from tip of snout and 
lower jaw to caudal base. Caudal spot vague or lacking. Distinct, usually unpig- 
mented stripe above black lateral stripe, about | scale row wide. Pre- and post- 
dorsal stripe narrow, about 3—5 melanophores wide; stripe slightly expanded at 
origin of dorsal fin. Lateral-line pores on anterior half of body outlined with 
melanophores. Scales above unpigmented lateral stripe with melanophores on 
posterior edge of scale pocket; free scale edge forming a posterior band on ex- 
posed field leaving anterior central portion unpigmented. Thin black line on pos- 
terior edge of scale giving double-banded appearance to each scale. Top of head 
dark and uniformly pigmented. Body unpigmented ventral to lateral line, except 


VOLUME 96, NUMBER | 55 


Table 3.—Counts of vertebrae (including the Weberian apparatus as four and the urostylar vertebra 
as one) in Notropis boops. East of Mississippi River = Wabash (INHS 2937), Cumberland (INHS 
78328, UT 44.249), Tennessee (SIUC 308, UT 44.709), and Mississippi (INHS 6047) drainages; west 
of Mississippi River = Missouri (INHS 80497), White (INHS 81072), Arkansas (INHS 81092), Oua- 
chita (INHS 81032), and Red (INHS 80736) drainages. 


Trunk Caudal Total 
Region 17 18 19 20 a 16 17 18 19 2 35 BO B37 3 n x 
East of Mississippi R. | 2 55 8 W9,2 2 Alby OX) iA 1 34 37 4 76 36.6 
West of Mississippi R. i 30 1 Ws3 sy 2D 5 16.8 3 Bil ils 49 36.2 


some melanophores outline base of anal fin. Rays of dorsal and caudal fins out- 
lined by melanophores. Rays of pectoral, pelvic, and anal fins occasionally out- 
lined with a few scattered melanophores. 

Numerous collections of breeding individuals from throughout the range of the 
species indicate that no breeding colors develop except for some creamy white 
in the membranes of the dorsal, caudal, and pectoral fins. Living individuals are 
generally greenish dorsally, silver in lateral aspect, and white ventrally. 

Tubercles well-developed in breeding males from April through August, poorly 
developed in females and non-breeding males. Peak of tubercle development 
appears to be in May. Males with large dense tubercles dorsal and posterior to 
eyes, on front of snout (below and anterior to nares), and mandibles. Top of head 
with minute tubercles densely distributed. Ventral surface of head, gill mem- 
branes, preopercle, and opercle with sparsely scattered small tubercles. Small 
tubercles outline scales above lateral line, being prominent anteriorly and dor- 
sally, and decreasing in size posteriorly and ventrally. Scales below lateral line 
and anterior to anal fin faintly outlined with tubercles. Small tubercles in single 
rows, mostly medially and distally, on branched and unbranched portions of all 
anal-fin rays, first 2 pelvic-fin rays, and first 5—7 dorsal-fin rays. Pectoral-fin tu- 
bercles on dorsal side of first 3-8 rays, occurring in double rows proximally and 
distally and densest (usually 4 rows) medially. Those on first unbranched ray 
rather evenly distributed in 2—4 rows on middle two-thirds of ray. Females rarely 
develop tubercles on chin; they are smaller and more sparsely distributed than 
those found on chin of males. 

Variation.—No significant sexual differences in meristic characters were noted. 
Excluding dimensions directly affected by gonadal development, males in breed- 
ing condition display significantly greater values for lengths of the anal, pectoral, 
and pelvic fins, and the preanal distance (see Student’s t-Tests, Table 1). 

The urogenital papilla of adult females is enlarged and protruded during the 
spawning season; the papilla of males is only slightly swollen. There are no 
significant sexual differences in pigmentation pattern. 

Notropis boops is remarkably uniform in external morphological features 
throughout its rather extensive range. Only two characters were found to exhibit 
some variation, the number of vertebrae and the number of lateral-line scales. 
Variation is slight in both characters, but the pattern of variation is similar. Spec- 
imens from east of the Mississippi River and in the northern part of the species’ 
range average slightly higher numbers of lateral-line scales and caudal vertebrae 
(Tables 2—3). The variation in these two characters follows a general trend re- 


56 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


KH 


j 200 300 


ol SCale OF Kilomerers | i 
Wiles 
f A S LZ prs is rai, eI 


Fig. 2. Range of N. boops. Solid dots represent localities from which specimens were examined; 
triangles represent localities from the literature; open circles represent localities where the species 
presumably no longer occurs. 


ported for many fishes in which the number of elements (e.g., vertebrae) pro- 
gressively increases to the north (Barlow 1961). 

Smith and Hocutt (1981) examined pharyngeal tooth counts in 100 individuals 
of N. boops from Big River, Missouri, and reported the following variation: 64% 
had a 1,4 4,1 count, 19% a 1,4-4,0 count, 8% a 0,4—4,0 count, 7% a 0,44, 1 
count, 1% a 1,4—3,1 count, and 1% a 1,5—4,1 count. Pharyngeal tooth counts 
from throughout the range of the species agree for the most part with those of 
Smith and Hocutt and indicate that a tooth is occasionally lost from the minor 
row. 

Comparisons.—Among the species of Notropis, N. boops most closely resem- 
bles N. xaenocephalus (Jordan) as Swift (1970) originally suggested. These two 
species are extremely similar in overall body shape and pigmentation pattern (Fig. 
1). Notropis xaenocephalus is, however, readily distinguished from N. boops by 
the following characters: pharyngeal teeth usually 2,4—4,2, anal-fin rays usually 
7, peritoneum silvery with scattered melanophores, caudal spot well-developed, 
and tuberculation on the head, body, and fins less extensive. 


VOLUME 96, NUMBER 1 57 


Distribution 


Notropis boops occurs in the central Mississippi basin where its range extends 
from the Scioto River drainage, Ohio, west to the lower Red River drainage, 
Oklahoma, south to the Ouachita River drainage, Louisiana, and north to the 
Illinois River drainage, Illinois (Fig. 2). Its reported occurrence in the Cache 
Creek drainage (Comanche County) of southwestern Oklahoma (Miller and Ro- 
bison 1973) is not verified by specimens. 

In recent years some populations of N. boops have been severely decimated, 
particularly in western Ohio (Trautman 1981) and east-central Illinois (Smith 1979) 
because of excessive siltation and turbidity. The continued use of poor agricul- 
tural practices in these states has modified many of the clear, gravel-bottomed 
streams that N. boops formerly inhabited (Fig. 2). 

The species is on the endangered or rare fish list for Mississippi (Clemmer ef 
al. 1975) and is of special concern in Alabama (Ramsey 1976). Although N. boops 
is rare or disappearing on the edges of its range it is common and abundant 
throughout most of Arkansas, eastern Oklahoma, southern Missouri, southeast- 
ern Kansas‘and central Kentucky and Tennessee. 


Acknowledgments 


We are grateful to the following ichthyologists and their institutions for infor- 
mation on types, loans of specimens, and/or providing one of us (BMB) with 
laboratory space: L. M. Page, Illinois Natural History Survey (INHS); F. B. 
Cross and J. T. Collins, University of Kansas (KU); K. Hartel, Museum of 
Comparative Zoology (MCZ); E. A. Lachner, U.S. National Museum of Natural 
History (USNM); T. M. Cavender, Ohio State University Museum of Zoology, 
(OSU); L. G. Hill, University of Oklahoma Museum of Zoology (OQUMZ); W. D. 
Pearson, University of Louisville (UL); R. M. Bailey, University of Michigan 
Museum of Zoology (UMMZ); D. A. Etnier, University of Tennessee (UT). K. 
Schmitt, Scientific Photography and Illustration Facility of the Southern Illinois 
University at Carbondale (SIUC) Graduate School, assisted in preparation of the 
figures. 


Literature Cited 


Barlow, G. W. 1961. Causes and significance of morphological variation in fishes.—Systematic 
Zoology 10(3):105—117. 

Clemmer, G. H., R. D. Suttkus, and J. S. Ramsey. 1975. A preliminary check-list of endangered 
and rare fishes of Mississippi, pp. 6-11. In A preliminary list of rare and threatened vertebrates 
in Mississippi.—Mississippi Game and Fish Commission, Jackson. 

Gilbert, C. H. 1884. A list of fishes collected in the East Fork of White River, Indiana, with 
descriptions of two new species.—Proceedings of the United States National Museum 7(423): 
199-205. 

Gilbert, C. R. 1978. Type catalogue of the North American cyprinid fish genus Notropis.—Bulletin 
of the Florida State Museum, Biological Sciences 23(1):1—104. 

Hubbs, C. L., and K. F. Lagler. 1964. Fishes of the Great Lakes region.—University of Michigan 
Press, Ann Arbor. 213 pp. 


58 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Kafuku, T. 1958. Speciation in cyprinid fishes on the basis of intestinal differentiation, with some 
references to that among catostomids.—Bulletin of the Freshwater Fisheries Research Labo- 
ratory 8:45-78. 

Lehtinen, S., and A. A. Echelle. 1979. Reproductive cycle of Notropis boops (Pisces: Cyprinidae) 
in Brier Creek, Marshall County, Oklahoma.—The American Midland Naturalist 102(2):237— 
243. 

Miller, R. J., and H. W. Robison. 1973. The fishes of Oklahoma.—Oklahoma State University 
Press, Stillwater. 246 pp. 

Ramsey, J.S. 1976. Freshwater fishes, pp. 53-65. Jn H. Boschung (ed.). Endangered and threatened 
plants and animals of Alabama.—Bulletin Alabama Museum of Natural History No. 2. 

Smith, P. W. 1979. The fishes of Illinois.—University of Illinois Press, Urbana. 314 pp. 

Smith, R. E., Jr., and C. H. Hocutt. 1981. Formulae variations of pharyngeal tooth counts in the 
cyprinid genus Notropis.—Copeia 1981(1):222—224. 

Snelson, F. F., Jr. 1971. Notropis mekistocholas, a new herbivorous cyprinid fish endemic to the 
Cape Fear River basin, North Carolina.—Copeia 1971(3):449—-462. 

Swift, C. C. 1970. A review of the eastern North American cyprinid fishes of the Notropis texanus 
species group (subgenus Alburnops), with a definition of the subgenus Hydrophlox, and ma- 
terials for a revision of the subgenus Alburnops.—Ph.D. thesis, Florida State University, 
Tallahassee. 515 pp. 

Trautman, M. B. 1981. The fishes of Ohio.—Ohio State University Press, Columbus. 782 pp. 


(BMB) Department of Zoology, Southern Illinois University at Carbondale, 
Carbondale, Ilinois 62901; (WWD) Museum of Zoology, Louisiana State Uni- 
versity, Baton Rouge, Louisiana 70893. 


PROC. BIOL. SOC. WASH. 
96(1), 1983, pp. 59-66 


SYNONYMY AND DISTRIBUTION OF 
PHYLLOMEDUSA BOLIVIANA BOULENGER 
(ANURA: HYLIDAE) 


David C. Cannatella 


Abstract.—YVhe correct name for the large, black-eyed species of Phyllomedusa 
from Bolivia, adjacent Brazil and northern Argentina is Phyllomedusa boliviana; 
P. pailona Shreve is a junior subjective synonym. The tadpole is a generalized 
phyllomedusine type. With the exception of the syntypes, none of the specimens 
referred to P. boliviana in the literature has been identified correctly. 


Boulenger (1902) described Phyllomedusa boliviana on the basis of a male and 
female from Chulumani, on the eastern slopes of the Andes in Bolivia. Until the 
revision by Funkhouser (1957), no additional specimens were referred to that 
species. Funkhouser did not examine the type specimens, and her relegation of 
several specimens from Buena Vista, Bolivia, to this species was based solely on 
the description of the species. 

In the course of studies on the systematics of phyllomedusine frogs, I have 
examined all of Funkhouser’s material and one of the syntypes of P. boliviana. 
None of the specimens examined by her can be ascribed to that species. Rather, 
they represent an undescribed species apparently related to P. tarsius, P. trini- 
tatis, and P. venusta, and will be treated in a paper dealing with the systematics 
of these species. 

In addition, I have collected P. boliviana at the type-locality and also in north- 
ern Argentina, where the species is known as Phyllomedusa pailona, described 
from El Pailon, Bolivia, by Shreve (1959). Live specimens from Chulumani and 
Argentina are similiar in the possession of a blackish-brown iris and eyelid edged 
in red. Direct comparison of the holotype and paratypes of P. pailona with the 
syntype of boliviana leads me to conclude that only one species is represented, 
and that Phyllomedusa boliviana is the proper name for the species. 


Materials and Methods 


Measurements were taken as described by Duellman (1970). The following 
abbreviations are used: snout—vent length, SVL; standard length (distance from 
tip of snout to tip of coccyx) SL; tibia length, TIB; foot length, FOOT; head 
length, HLEN; head width, HWID; interorbital distance, IOD; internarial dis- 
tance, IND; length of eyelid, ELID; length of eye, EYE; horizontal diameter of 
tympanum, TY MP; distance from anterior corner of eye to nostril, ENOS. Color 
notes from life are taken from the field notes of the author. The following museum 
acronyms were employed: American Museum of Natural History, AMNH; Brit- 
ish Museum (Natural History), BMNH; Carnegie Museum, CM; Instituto Miguel 
Lillo (Tucuman, Argentina), IML; University of Kansas Museum of Natural His- 
tory, KU; Museo Argentino de Ciencias Naturales, (Buenos Aires, Argentina), 
MACN; Museum of Comparative Zoology, MCZ; University of Michigan Mu- 


60 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


seum of Zoology, UMMZ; United States National Museum of Natural History, 
USNM. 


Phyllomedusa boliviana Boulenger 
Fig. | 


Phyllomedusa boliviana Boulenger, 1902:395. Syntypes, BMNH 1901.8.2.49-S0 
(RR 1947.2.22.32-33) from Chulumani, Bolivia, 2000 m, P. O. Simons, collec- 
tor. 

Phyllomedusa (Pithecopus) boliviana.—Lutz, 1950:602, 621. 

Phyllomedusa pailona Shreve, 1959:1. Holotype, MCZ 29677, El Pailon, 5 km 
from the eastern shore of the Rio Grande, Depto. Santa Cruz, Bolivia, 350 m, 
Carl Gans and F. S. Pereira, collectors.—Duellman, 1968:6.—Duellman, 1977: 
163. 

Pithecopus pailonus.—Lutz, 1966:236.—Laurent, 1967:232. 


Diagnosis.—A large species of Phyllomedusa having 1) the first toe longer than, 
and opposable to, the second; 2) distinct, well-developed parotoid glands; 3) 
prevomerine teeth present in adults; 4) no calcar or dermal appendages; 5) a 
lightly reticulated palpebral membrane; 6) iris black-brown in life; 7) in life, upper 
eyelid edged with red, tubercles on flank white, tipped with red; concealed sur- 
faces of hindlimb pale yellow-green with no markings. 

Redescription of male syntype.—Head wider than body; snout short, rounded 
in dorsal view; in lateral view, truncate and inclined posteriorly from lip to nostril; 
canthus rostralis rounded, distinct; loreal region barely concave; lips thin and not 
flared; nostrils not protuberant, directed laterally; internarial region flat; eyes not 
protuberant; pupil vertically elliptical; palpebrum finely reticulated; parotoid gland 
well-developed, extending from eyelid to a point level with the insertion of the 
arm; supratympanic fold moderately developed, barely obscuring upper edge of 
tympanum; tympanum distinct, oval; anterior edge of tympanum separated from 
posterior corner of eye by distance of 1 mm. Axillary membrane absent; upper 
arm slender, forearm robust; ulnar fold low; relative lengths of fingers 1-2-4-3 
from shortest to longest; finger discs about three-fifths diameter of tympanum; 
subarticular tubercles large, round to conical; supernumerary tubercles lacking 
on fingers; palmar tubercle barely distinct; prepollex enlarged, elliptical, bearing 
thin, horny nuptial excrescence; fingers lacking webbing. 

Leg of moderate length, slender; no calcar or dermal ornamentation; inner 
tarsal fold absent; outer tarsal fold barely distinct; relative lengths of toes 2-1-3- 
5-4 from shortest to longest; toe discs rounded, shorter than those on fingers; 
inner metatarsal tubercle elliptical, low, flattened; outer metatarsal tubercle ab- 
sent; no webbing between toes; subarticular tubercles large, round to conical; no 
supernumerary tubercles on toes. 

Anal opening directed posteriorly at midlevel of thighs; anal opening a short 
tube, no flap; supra-anal fold present; pair of large tubercles present inferior and 
lateral to anus; skin of dorsal surfaces smooth, no tubercles; skin of chin and 
pectoral region finely areolate; skin of belly and ventral surface of thigh tuber- 
cular; skin of remaining ventral surfaces of limbs smooth, except for that of 
forearm and tarsus, which bears a few discrete tubercles; anterior flank tuber- 
culate, posterior flank smooth; tongue lanceolate, barely notched posteriorly; free 


VOLUME 9%, NUMBER 1 61 


Fig. |. Phyllomedusa boliviana, male, KU 182969, SVL 64.8 mm. 


for about one-half its length; prevomerine teeth present; dentigerous processes 
of prevomer small, separated medially by distance equal to width of one process, 
processes directed posteromedially at midlevel of elliptical choana; vocal slit 
short, extending from posterolateral corner of tongue to corner of mouth; vocal 
sac single, median, subgular. 

In preservative, dorsal surfaces pale blue; dorsal surfaces of finger and toe 
discs gray; ventral surface of chin pale brown, with few diffuse cream spots; 
ventral surface of belly and thigh cream; ventral aspects of forearm and tarsus 
pale brown, with discrete white tubercles; ventral surfaces of hand and foot pale 
brown; ventral surfaces of shank and inner aspect of tarsus pale gray with pale 
brown mottling; anterior and posterior aspect of thigh pale blue-violet; few, in- 
distinct white tubercles present on posterior aspect of thigh; anal region pale 
brown; paired anal tubercles and supra-anal fold white; white stripe present along 
heel, tarsus, and lateral border of fifth toe; border of upper eyelid, lower lip, and 
anterior flank white; most tubercles of anterior flank capped with small spot of 
dark gray pigment; white stripe at point of insertion of arm, demarcating the blue 
color of upper arm from cream region of chest; this stripe and lip stripe bordered 
inferiorly by thin gray line. 

The male syntype is in excellent condition, except that the outer two toes of 
each foot are slightly desiccated. Also, there is a | cm incision in the midventral 
region. The female specimen is similar to the male syntype in most respects; 
Duellman (pers. comm.), however, noted that the palpebrum of the female was 
not reticulated. 

Measurements of the syntypes (male/female) in mm.—SVL 54.3/75.2, TIB 23.9/ 


62 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 2. Distribution of P. boliviana. Key to localities: 1. Caranavi; 2. Chulumani; 3. El Pailon; 4. 
Jaciara; 5. Aguas Blancas, Rio Pescado, and Finca Jakulica; 6. Yuto. 


1, FOOT 18.0/23.7, HLEN 18.8/22.0, HWID 18.6/23.8, IOD 5.2/8.1, IND 4.1/ 
Sel EID YAS 25.7 ENE 691/823) NOMIR 322/42 ENO Sis20/5222 

Coloration in life.—Dorsal surfaces, face, and tympanum lime green; upper 
eyelid edged with red; venter gray, with pale yellow splotches; lower lip, supra- 
anal fold, and tarsal fold white, edged with pale red; tubercles on flank white, 
tipped with red; tarsal fold white; ventral surfaces of hand and foot flesh-colored; 
axilla and concealed region of elbow pale yellow; groin, concealed surface of 
shank, tarsus, and thigh pale yellow-green with a hint of orange; anal region dark 
gray; paired anal tubercles pale yellow; finger and toe discs off-white; iris dark 
brown, almost black (D. Cannatella field notes, 12 December 1978). 

The above notes are based on a sample of Argentinian specimens (KU 182969— 
90). Specimens from Chulumani, Bolivia (KU 183436—38) differed in the following 
points: the tubercles on the ventral surface of the forearm and tarsus, and the 
supra-anal fold were eee the ulnar and tarsal stripes were white with a 
red-orange wash. 

Distribution.—This species is known from the Andean slopes and eastern low- 
lands of Bolivia, and the lowlands of western Brazil and northern Argentina, at 
elevations of 350-2000 m (Fig. 2). Duellman (1977) followed Funkhouser (1957) 
in including Peru within the species’ range, on the basis of her misidentified 
specimens. Likewise, the specimens from Restauragao, Brazil, on which Heyer 
(1977) remarked, were compared to misidentified “‘boliviana’’; the frogs from 
Restauracgao are not boliviana, but are referrable to the undescribed species from 
Buena Vista mentioned above. 

Variation.—The syntypes of P. boliviana and the type-series of P. pailona 


VOLUME 96, NUMBER 1 63 


Table 1.—Measurements (mm) of P. boliviana from northern Argentina; mean + 2 standard errors: 
range in parentheses. 


nnn, tee agg 


Measurement Males (n = 19) Females (n = 3) 
SVL 67.7 + 1.18 76.8 + 3.54 
(63.6—74.2) (74.8-80.3) 
SL 65.3 + 1.18 72.3 + 3.10 
(60.5—70.8) (69.9-75.2) 
TIB 28.4 + 0.47 31.4 + 0.59 
(27.1-30.9) (30.8—-31.8) 
FOOT 23.1 + 0.49 Mol) = NIKO 
(20.8—24.6) (24.4-26.7) 
HLEN 23.2 = 0.28 24.6 + 1.40 
(22.0-24.6) (23.9-26.0) 
HWID 23.0 + 0.28 24.6 + 0.54 
(22.2-24.3) (24.2—25.1) 
IND 5.4 + 0.12 5.9 + 0.43 
(4.9-6.0) (5.5—6.2) 
ENOS 5.4 + 0.06 6.0 = 0.30 
(5.25.6) (5.7-6.2) 
TYMP 4.1 + 0.13 4.5 + 0.20 
(3.64.4) (4.44.7) 


differ in ventral coloration. The syntypes of P. boliviana and the KU specimens 
from Chulumani and Caranavi, Bolivia, have flank tubercles tipped with dark 
brown spots; the venters are dark brown with areas of pale mottling (some have 
no pale areas). Also, the venter of the male syntype of P. boliviana is slightly 
paler than those of the KU specimens. 

One of the specimens of the type-series of P. pailona has just a hint of such 
flank spots (CM 36278) and the others lack them. Moreover, two of these have 
a white lateral stripe on the anterior flanks, bordered below by a dark blue stripe. 
The venters of these frogs are almost immaculate, a diagnostic feature noted by 
Shreve (1959). 

Most of the frogs from Jaciara, Brazil, have a hint of the dark tips on the flank 
tubercles, and all have a more discrete lateral stripe as well, bordered below by 
a dark blue-gray stripe or wider area. The venters of this series are pale gray, 
slightly darker than the type-series of pailona, but not as dark as the Chulumani 
series. 

The frogs from Agua Blanca, Argentina, consist of two series: one recently 
collected by me and the other collected by Dr. Raymond Laurent. The specimens 
collected by me have dark brown venters with pale mottling, and a few discrete 
dark spots on the flank tubercles. The older specimens have immaculate venters 
with almost no hint of the flank tubercle spots. The KU specimens from Finca 
Jakulica and Angosto del Pescado, Argentina are colored similarly to the recent 
KU specimens from Agua Blanca. 

It appears that the striking differences in ventral coloration between the type- 
specimens of both nominal species are due to some preservation difference and/ 
or fading over time. Additionally, all of the recently collected KU specimens 
have dark blue dorsal surfaces, and those of the other specimens are pale blue. 


64 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 3. Larva of Phyllomedusa boliviana, KU 183822. 


In the preserved specimens there is also variation in the presence of a reticu- 
lated palpebrum, but this is clearly a preservation artifact. All of the specimens 
collected by me (KU 182969-90 and KU 183436—40) possessed a reticulated pal- 
pebrum in life. However, these specimens were stored for several months in 
formalin before transfer to ethanol, and the reticulation has been lost in all of 
them. All other specimens that I have examined still possess the reticulation. 

Measurements of the specimens collected in northern Argentina are summa- 
rized in Table 1. Males from the vicinity of Chulumani, Bolivia range from 54.3 
to 70.2 mm SVL (n = 3). The two males from Caranavi are 53.5 and 57.2 mm 
SVL. Those from El Pailon, Bolivia (n = 4) range from 56.1 to 64.7 mm SVL, 
and the males from Jaciara are 62.1—68.3 mm in SVL (n = 6). 

Justification of synonymy.—Direct comparison of the type-specimens of P. 
boliviana and P. pailona leaves little doubt that the two names have been applied 
to the same species. Shreve based his comparison on material of “‘boliviana”’ 
that had been misidentified by Funkhouser (1957). As noted above, the immac- 
ulate venter of P. pailona is due to preservation artifacts. The iris color alone 
will diagnose P. boliviana from all other Phyllomedusa. Additionally, the com- 
bination of reticulated palpebrum, large parotoid glands, and absence of pattern 
on the posterior flanks and concealed surfaces of the hindlimbs will distinguish 
this species from all others of the genus. 

Natural history.—At Aguas Blancas, Argentina, specimens were collected 0.5— 
1.0 m above ground in vegetation near ruts filled with water at the side of a 
highway. The males were calling; the call is a ‘tup-tup-tup,’ repeated every 2-10 
seconds. At Angosto del Pescado and Finca Jakulica, individuals were collected 
around ponds or ditches filled with water, about 1 m above ground in vegetation; 
the males were calling in the same fashion. Barrio (1976) described the mating 
call of this species from the Rio Pescado, Argentina. 

In Bolivia, individuals were calling from vegetation about 1 m above ground; 
none were collected near standing water, however. All specimens collected by 
me were taken at night. Laurent (1967) remarked on the natural history of the 
species in Argentina. 

Tadpoles.—The lot of tadpoles was collected by day from a muddy roadside 
ditch, where KU 18343940 were taken on the previous night. Tadpoles of Ololy- 
gon rubra were collected in the same ditch. 

In general, the larvae of P. boliviana resemble those phyllomedusine larvae 
described by Cannatella (1980, 1982). The following description is based on KU 
183822, which consists of seven tadpoles. A representative larva at Stage 32 


VOLUME 96, NUMBER 1 65 


(Gosner 1960) with a SVL of 15.0 mm and total length of 42.5 mm is illustrated 
in Fig. 3. Body slightly deeper than wide, deepest and widest at two-thirds the 
length of the body; top of head slightly convex; snout rounded in lateral profile; 
snout rounded in dorsal view; nostrils dorsolateral; internarial distance slightly 
greater than width of oral disc; eyes dorsolateral and directed laterally; spiracle 
a flap-like tube, ventral and sinistral to midline; spiracular opening at a point 
about midlength of the body; mouth anteroventral; cloacal tube short, dextral to 
base of caudal fin; caudal musculature slender, tapering gradually to posterior 
end of fin; myomeres weakly developed; at midlength of the tail the depth of the 
caudal musculature slightly less than depth of ventral fin, but greater than depth 
of dorsal fin; caudal musculature extending to tip of tail; dorsal fin shallow an- 
teriorly, not extending onto body; dorsal fin deepest at two-thirds its length from 
anterior; ventral fin deepest at its midlength. 

Mouth relatively small, with a shallow lateral fold; medial portion of upper lip 
lacking papillae; elsewhere, papillae present in one or two rows along borders; 
upper beak broadly curved and uniformly serrate; lower beak V-shaped and uni- 
formly serrate; two upper and three lower rows of denticles; upper rows of same 
length; second upper row broadly interrupted medially; three lower rows of den- 
ticles uninterrupted; first two lower rows about same length; third row about one- 
half as long as upper two; denticles of third row smaller and fewer in number. 

In life, dorsum transparent with gold and black pigments; lateral surfaces gold; 
venter transparent, with some gold pigment anteriorly; tail clear with fine distri- 
bution of melanophores; area above eye heavily pigmented with gold; iris white 
with black and gold flecks. In preservative, sides and top of head and body 
translucent, with scattered melanophores; intense concentration of melanophores 
covering gut, and obscuring view of intestines; caudal musculature flesh-colored; 
dorsal and ventral fins transparent, with fine distribution of melanophores in 
posterior one-half of dorsal and ventral fins, but not extending to tip of tail. 

Eggs.—Three clutches of eggs were examined (IML 1349). One clutch is a lot 
of hatchling tadpoles. The second is a clutch of 154 tadpoles about to hatch from 
the egg membranes; the egg mass is still enveloped in a folded leaf. The third 
clutch is also encased in a folded leaf and consists of 142 eggs in the first stages 
of cleavage; the mean diameter of 20 eggs is 2.6 mm. A few of the eggs appear 
to be unfertilized and are smaller in size, and empty egg cases are found at the 
upper and lower ends of the cylindrical clutch in the same fashion as described 
for P. hypocondrialis by Pyburn (1980), and P. duellmani by Cannatella (1982). 

Specimens examined.—ARGENTINA: Jujuy: Ruta Yuto-Ledesma, a 7 km de 
bifurcacion, IML 1305(7), 1307(8); Bifurcacion ruta Yuto, 1306(2), 1308; Pozo 
Colque (cerca de Yuto), 1309(3); Alrededores de Yuto, 1310; Yuto, UMMZ 127406; 
Salta: Rio Pescado, IML 1345(24), 1397(8), 1349 (eggs); Aguaray, 1467(6); Abra 
Grande, Oran, 1682(2); Agua Blanca, 460 m, 1685 (11), 2145 (4), 2171, KU 182969- 
71, KU 12894044, 128945 (skeleton); Angosto del Pescado 620 m, IML 2706(24), 
1349(3) (three clutches eggs), KU 182972-75; Finca Jakulica, 560 m, 182976—-90. 

BRAZIL: Mato Grosso: Jaciara, UMMZ 127908, USNM 164097, MCZ 44942, 
AMNH 72450, KU 92322-25, 92326—27 (skeletons). 

BOLIVIA: La Paz: 4.4 km E Chulumani, 1760 m, KU 183436; 2.1 km E Chu- 
lumani, 1900 m, 183437-38; Chulumani, 2000 m, BMNH 1947.2.22.33 (syntype); 
15.7 km S Caranavi, 900 m, KU 183439-40, 183822 (larvae); Santa Cruz: Rio 


66 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Grande, Cabezas, MACN 3483(5); El Pailon, 350 m, CM 36278-79 and MCZ 
29679 (paratypes of P. pailona); MCZ 29677 (holotype of P. pailona). 


Acknowledgments 


For loan of specimens and provision of working space I thank Charles W. 
Myers, Alice G. C. Grandison, Jose M. Gallardo, Raymond F. Laurent, Arnold 
G. Kluge, W. Ronald Heyer, C. J. McCoy, and Pere Alberch. Raymond Laurent 
was a gracious host during my stay in Argentina, and Esteban LaVilla and Omar 
Pagaburo greatly assisted me there in the field. Thomas Berger travelled with me 
in Bolivia. William E. Duellman shared data on the female syntype of P. boliviana 
and also criticized the manuscript. Collecting permits for Bolivia and Argentina 
were granted by Gaston Bejarano B. of the Departamento Nacional de Vida 
Silvestre, Parques Nacionales, Caza y Pesca, of Bolivia and Malcolm D. Craig, 
Direccion Nacional de Fauna Silvestre, Argentina. Field work was supported by 
a grant (DEB 76-09986, William E. Duellman, principal investigator) from the 
National Science Foundation, and by a National Science Foundation Graduate 
Fellowship to the author. 


Literature Cited 


Barrio, A. 1976. Estudio cariotipico y analisis audioespectrografico de los cantos de las especies 
de Phyllomedusa (Anura, Hylidae) que habitan en la Argentina.—Physis 35(90):65—74. 
Boulenger, G. A. 1902. Descriptions of new batrachians and reptiles from the Andes of Peru and 

Bolivia.—Annals and Magazine of Natural History (7)10:394—402. 

Cannatella, D. C. 1980. A review of the Phyllomedusa buckleyi group (Anura: Hylidae).—Occa- 
sional Papers of the Museum of Natural History of the University of Kansas (87):1—40. 
——. 1982. Leaf-frogs of the Phyllomedusa perinesos group (Anura: Hylidae).—Copeia 1982(3): 

501-513. 

Duellman, W. E. 1968. The genera of phyllomedusine frogs (Anura: Hylidae).—University of Kan- 
sas Publications, Museum of Natural History 18(1):1—10. 

——. 1970. The hylid frogs of Middle America.—Monograph of the Museum of Natural History, 
University of Kansas (1):1—753. 

———. 1977. Liste der rezenten Amphibien und Reptilien: Hylidae, Centrolenidae, Pseudidae.— 
Das Tierreich (95): 1—225. 

Funkhouser, A. 1957. A review of the Neotropical tree-frogs of the genus Phyllomedusa.—Occa- 
sional Papers of the Natural History Museum of Stanford University (5): 1—90. 

Gosner, K. L. 1960. A simplified table for staging anuran embryos and larvae with notes on iden- 
tification.—Herpetologica 16(3): 183-190. 

Heyer, W. R. 1977. Taxonomic notes on frogs from the Madeira and Purus Rivers, Brasil.—Papéis 
Avulsos de Zoologia, Sao Paulo 31(8):141—162. 

Laurent, R. F. 1967. Redescubrimiento de Pithecopus pailonus (Shreve) en Argentina.—Acta Zoo- 
logica Lilloana 22:231—248. 

Lutz, B. 1950. Anfibios anuros da colécao Adolpho Lutz. Hylidae in the Adolpho Lutz collection 
of the Instituto Oswaldo Cruz.—Memorias do Instituto Oswaldo Cruz 11(3):599-637. 

——. 1966. Pithecopus ayeaye, a new Brazilian hylid with vertical pupils and grasping feet.— 
Copeia 1966(2):236—-240. 

Pyburn, W. F. 1980. The function of eggless capsules and leaf in nests of the frog Phyllomedusa 
hypochondrialis (Anura: Hylidae).—Proceedings of the Biological Society of Washington 93(1): 
153-167. 

Shreve, B. 1959. A new Phyllomedusa from Bolivia (Salientia, Hylidae) —Breviora (113):1-3. 


Museum of Natural History and Department of Systematics and Ecology, The 
University of Kansas, Lawrence, Kansas 66045. 


PROC. BIOL. SOC. WASH. 
96(1), 1983, pp. 67-78 


STUDIES OF THE COASTAL MARINE FAUNA OF 
SOUTHERN SINALOA, MEXICO. IV. REPORT 
ON THE CARIDEAN CRUSTACEANS 


M. E. Hendrickx, M. K. Wicksten, and A. M. van der Heiden 


Abstract.—As a result of a large scale survey of the coastal marine fauna of 
southern Sinaloa, Mexico, 29 species of caridean shrimps are reported for a wide 
variety of habitats. Information is provided on their currently known distribution 
in the Eastern Pacific region. 


Since 1979, a large scale survey of the marine coastal fauna has been underway 
in the southeastern part of the Gulf of California. One of the main results obtained 
so far is the updating of a provisional inventory of the species of marine and 
estuarine invertebrates and fishes found in southern Sinaloa (van der Heiden and 
Hendrickx 1982; Hendrickx and van der Heiden in press). Much information also 
has been gained regarding the occurrence of these species in different habitats. 

There have been almost no faunistic studies on invertebrates on a regional 
basis in the Gulf of California. The most comprehensive works available are those 
of Parker (1963) and Brusca (1980). Both, however, are incomplete, mentioning 
only some of the wide variety of species to be found in the area. This paper 
records the caridean shrimps found so far in southern Sinaloa, and provides 
information on their habitats. Further studies are underway, however, and ad- 
ditional species may be found to be present in the area, especially in less acces- 
sible habitats such as the nearshore rocky subtidal. 

Material and methods.—TYhe material on which this study of caridean shrimps 
is based comes from a wide spectrum of habitats with differing environmental 
conditions. Specimens from shallow water coastal systems came from estuaries, 
coastal lagoons, and rocky intertidal habitats. Material from the Bay of Mazatlan 
was taken during a 2-year sampling program aboard the boat FC-/ of the Secre- 
taria de Educacion Publica in Mazatlan. Specimens from the continental shelf 
were collected during a three-leg project (SIPCO project) aboard the oceano- 
graphic vessel E/ Puma of the Instituto de Ciencias del Mar y Limnologia, Uni- 
versidad Nacional Autonoma de México. Sampling in shallow water was done 
by hand collecting, cast nets, trawls, small grabs, or small dredges. Aboard ship, 
bottom grabs, dredges, beam trawls, or otter trawls were used according to the 
capabilities of the ships that were available. A map of the sampling areas is given 
in Fig. 1. 


Family Pasiphaeidae 
Leptochela serratorbita Bate, 1888 


Material.—Bay of Mazatlan, Van Veen grab: 2 males (April 1980), | ovigerous 
female (Jan 1980). 

Previous Eastern Pacific records.—Cape San Lucas, Baja California Sur; To- 
polobampo, Sinaloa (Chace 1937). 


68 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Range.—Tropical and warm temperate western Atlantic (Chace 1972), southern 
Gulf of California. 

Habitat.—At Cape San Lucas, 15-30 m on sandy bottom (Chace 1937); in Bay 
of Mazatlan, 10-27 m on fine sand. 


Family Palaemonidae 
Palaemon (Palaemon) gracilis (Smith, 1871) 


Material.—Caimanero lagoon, cast net, 23 specimens (June 1978; Sept 1978; 4 
ovigerous). 

Previous Eastern Pacific records.—Nicaragua and Panama (Holthuis 1952b). 

Range.—Caimanero lagoon south to Rio Lara, southern Panama. 

Habitat.—Rivers, streams and coastal waterways throughout its range, in fresh 
to brackish water (up to 8%c salinity). 


Macrobrachium americanum Bate, 1868 


Material.—Rio Baluarte, 4 specimens (1979). 

Previous Eastern Pacific records.—Numerous records, from Mexico to Peru 
(Holthuis 1952b). 

Range.—Mulege, Baja California Sur to northern Peru (Holthuis 1952a). 

Habitat.—Freshwater streams in southern Sinaloa (Rio Presidio, Rio Baluarte, 
and Rio Quelite). 


Macrobrachium digueti (Bouvier, 1895) 


Material.—Rio Baluarte, 3 specimens (Nov 1978, Aug 1980). 

Previous Eastern Pacific records.—Baja California, Mexico; Guatemala, Pan- 
ama, and Ecuador (Holthuis 1952b); Peru (Méndez 1981). 

Range.—Mulege, Baja California Sur; Rio Baluarte, Sinaloa south to Rio Moche, 
Peru. 

Habitat.—Fresh water. 


Macrobrachium occidentale Holthuis, 1950 


Material.—Rio Baluarte, 2 specimens (Nov 1978). 

Previous Eastern Pacific records.—Guatemala, El] Salvador and Panama (Hol- 
thuis 1952b). 

Range.—Rio Baluarte, Sinaloa south to Panama. 

Habitat.—Specimens from the Rio Baluarte were collected in fresh water. 


Macrobrachium tenellum (Smith, 1869) 


Material.—Huizache-Caimanero lagoon complex, 3 specimens (July 1978); Cai- 
manero lagoon, cast net, 7 ovigerous females (Sept 1978); Estero El Verde, hand 
net, 3 specimens (July 1979). 

Previous Eastern Pacific records.—Numerous records, Mexico to Peru (Hol- 
thuis 1952b). 

Range.—Mulege, Baja California Sur to northern Peru (Holthuis 1952a). 

Habitat.—So far, the species has been reported only in fresh water. The ani- 


VOLUME 96, NUMBER 1 69 


PUNTA 
PIAXTLA 


ESTERO ee A 
EL VERDE ——> 


PUNTA LOS 
CERRITOS 


SEE MAP C 


BAY OF 
MAZATLAN 


ESTERO 
DE URIAS 


HULZACHE 
20 km LAGOON 


MAP B: REGION OF 


SOUTHERN SINALOA CAIMANERO 
LAGOON 


SEE pe DD, OM 2. ESTERO 
PAJAROS I. ie wig, oN NEG): DE TEACAPAN 


ba) cE ae Ie 
0 1 km ¥ 


——) VI 
MAP C: BAY OF MAZATLAN 


Fig. 1. Location of the sampling areas. Map A: Gulf of California; Map B: Region of Southern 
Sinaloa; Map C: Bay of Mazatlan. 


mals from near Mazatlan, however, were collected in brackish water. The salinity 
ranged up to 20%c. In Estero El Verde, M. tenellum was found in beds of widgeon 
grass (Ruppia sp.). 


Palaemonetes (Palaemonetes) hiltoni Schmitt, 1921 


Material.—Estero El Verde, 11 specimens, 9 ovigerous (June and Dec 1979; 
Jan and May 1980); Caimanero lagoon, cast net, 61 specimens (date not recorded). 

Previous Eastern Pacific records.—Southern California, U.S.A., Sonora and 
Sinaloa, Mexico (Holthuis 1952b). 

Range.—San Pedro, California south to Sinaloa. 


70 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Habitat.—Animals from estero El Verde were taken among widgeon grass 
(Ruppia sp.) with a small hand net at 0.4-1 m at a salinity of 20-22%o. 


Periclimenes (Periclimenes) infraspinis (Rathbun, 1902) 


Material.—Bay of Mazatlan, Van Veen grab, 3 specimens (Aug 1979). 

Previous Eastern Pacific records.—Southern California, U.S.A.; Gulf of Cal- 
ifornia, Costa Rica, Galapagos Islands (Holthuis 1951). 

Range.—San Diego, U.S.A., south to the Galapagos Islands (Brusca 1980). 

Habitat.—The specimens were taken on a sandy bottom at 9 m. 


Neopontonides dentiger Holthuis, 1951 


Material.—Bay of Mazatlan, trawl, 1 ovigerous female (Sept 1979); off Punta 
Piaxtla (23°34'N, 106°57’W), trawl, 1 specimen (Apr 1981). 

Previous Eastern Pacific record.—Off Cape San Francisco, Ecuador (Holthuis 
1951, 1952a). 

Range.—The species is known only from these 3 records. 

Habitat.—Holthuis (1951) reported the holotype from a mud-rock bottom at 4 
m. Material from Sinaloa was found a sand-rock bottom at 8 m in the Bay of 
Mazatlan, and on a muddy bottom with colonies of gorgonians attached to stones 
at 66 m off Punta Piaxtla. 


Pontonia margarita Smith, 1869 


Material.—Bay of Mazatlan: | male and 1 female (Sept 1979), 2 males and 2 
ovigerous females (June 1980). 

Previous Eastern Pacific records.—Numerous records, from the Gulf of Cali- 
fornia to the Galapagos Islands (Holthuis 1951). 

Range.—From the Gulf of California to the Galapagos Islands, Ecuador (Hol- 
thuis 1951). 

Habitat.—Specimens were found as commensals of the pearl oyster Pinctada 
mazatlanica. 


Family Gnathophyllidae 
Gnathophyllum panamense Faxon, 1893 


Material.—Punta Los Cerritos, north of Mazatlan, rocky shore, | specimen 
(May 1981). 

Previous Eastern Pacific records.—Gulf of California south to Panama (Brusca 
1980). 

Range.—Gulf of California south to the Galapagos Islands (Wicksten in press). 

Habitat.—Rocky intertidal zone to 17 m, sometimes found in tidepools. 


Family Hippolytidae 
Lysmata californica (Stimpson, 1866) 


Material.—Bay of Mazatlan, trawled, 3 specimens, | ovigerous (Nov 1979). 
Previous Eastern Pacific records.—California, U.S.A.; west coast of Baja Cal- 


VOLUME 96, NUMBER 1 71 


ifornia, Gulf of California, to Panama (Abele and Patton 1976; Brusca 1980; Stand- 
ing 1981). 

Range.—TYomales Bay, California to Panama. 

Habitat.—Common in rocky intertidal areas (Brusca 1980). Taken at 9 and 24 
m on muddy sand or fine sand off Mazatlan. 


Trachycaris restrictus (A. Milne-Edwards, 1878) 


Material.—Off Punta Piaxtla, trawled, 1 ovigerous female (Apr 1981). 

Previous Eastern Pacific records.—Gulf of California, Panama (Wicksten in 
press). 

Range.—Tropical eastern and western Atlantic (Chace 1972); Gulf of California 
to Panama. 

Habitat.—The specimen from off Punta Piaxtla was taken on a muddy bottom. 
at 66 m. 


Thor paschalis (Heller, 1862) 


Material.—Bay of Mazatlan, Van Veen grab, 1 specimen (Nov 1979); Pichi- 
lingue, Bay of La Paz, Baja California, rocky shore, 3 specimens (Mar 1980). 

Previous Eastern Pacific records.—Gulf of California, southwestern Mexico, 
Panama (Wicksten in press). 

Range.—Indo-West Pacific region (Bruce 1976); Gulf of California to Panama. 

Habitat.—At La Paz, the animals were found under stones in a rocky intertidal 
area. The animal from the Bay of Mazatlan was collected on a bottom of stones 
and shell fragments. 


Latreutes antiborealis Holthuis, 1952c 


Material.—Bay of Mazatlan, Van Veen grab, | specimen (Nov 1980). 

Previous Eastern Pacific records.—Numerous records, Gulf of California to 
Chile (Wicksten and Méndez in press). 

Range.—Gulf of California to Chile. 

Habitat.—The specimen from the Bay of Mazatlan was collected on fine to 
very fine sand at 6 m. 


Family Processidae 
Processa spp. 


Material.—Off Teacapan, Sinaloa (22°24’N, 105°54’W), Van Veen grab, | ovi- 
gerous female (Aug 1980); Bay of Mazatlan, Van Veen grab, 24 specimens from 
12 different grab samples (Aug 1979—Mar 1981). 

Habitat.—The animal from off Teacapan was taken at 37 m on silty sand; the 
rest came from 7-25 m on sandy mud to muddy sand, sometimes with shell 
fragments. 

Remarks.—Individuals of this genus are common in the Bay of Mazatlan. Most 
of these specimens belong to an undescribed species being studied by M. K. 
Wicksten and M. Méndez. Others have not yet been identified to species. 


72 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Family Alpheidae 
Synalpheus digueti Coutiere, 1909 


Material.—Pajaros Island, Bay of Mazatlan, rocky intertidal zone, 1 specimen 
(Apr 1980); Punta Chile, Bay of Mazatlan, rocky intertidal zone, 3 specimens 
(Oct 1980). 

Previous Eastern Pacific records.—**Lower California’ (Coutiére 1909); Arena 
Bank, Gulf of California (Chace 1937); Panama (Abele and Patton 1976); Isla 
Malpelo, Colombia (Abele 1975). 

Range.—Southern Gulf of California to Colombia. 

Habitat.—Lower midlittoral to sublittoral zones, among stones or within crev- 
ices in rocky intertidal; also associated with sponges in lower intertidal zone. 


Synalpheus nobilii Coutiere, 1909 


Material.—About 5 km N of Mazatlan, 2 specimens (1 ovigerous) (Dec 1979); 
Punta Chile, Bay of Mazatlan, 1 ovigerous female (Oct 1980); foot of Cerro de 
Vigia, Bay of Mazatlan, 2 specimens (1 ovigerous) (Oct 1980); Cerro de Vigia, 5 
specimens (2 ovigerous) (Nov 1980). 

Previous Eastern Pacific records.—Gulf of California (Wicksten in press); Clip- 
perton Island (Chace 1962); Panama (Abele 1976); Isla Malpelo, Colombia (Abele 
1975); Ecuador (Coutiére 1909); Galapagos Islands (Schmitt 1939). 

Range.—Gulf of California to the Galapagos Islands. 

Habitat.—All specimens except the female from Punta Chile came from be- 
neath tufts of coralline algae in the lower midlittoral zone of rocky shores. 


Synalpheus biunguiculatus (Stimpson, 1860) 


Material.—Punta Piaxtla, rocky shore, 2 specimens (1 ovigerous) (Oct 1979); 
Punta Chile, Bay of Mazatlan, rocky shore, 2 ovigerous females (Oct 1980); foot 
of Cerro de Vigia, Bay of Mazatlan, 5 specimens (1 ovigerous) (Oct 1980); Estero 
de Teacapan, 2 specimens (Mar 1980), 4 specimens (Apr 1980), 1 specimen (May 
1980). 

Previous Eastern Pacific records.—Gulf of California, southwestern Mexico, 
and Galapagos Islands (Wicksten in press); Clipperton Island (Chace 1962), Pan- 
ama (Abele 1976), Isla Malpelo, Colombia (Abele 1975). 

Range.—Hawaiian Islands (Stimpson 1860; Banner 1953), Gulf of California 
south to Galapagos Islands. 

Habitat.—Specimens from Punta Piaxtla and the Bay of Mazatlan were found 
under sponges on rocks and under stones. The animals from Estero de Teacapan 
were collected on oyster racks with Striostrea iridescens and barnacles (Balanus 
sp.) at a salinity of 40%c in March 1980). 


Synalpheus apioceros sanjosei Coutiere, 1909 


Material.—Punta Piaxtla, rocky shore, 1 ovigerous female (Oct 1980); Bay of 
Mazatlan, | specimen (June 1979); Punta Chile, Bay of Mazatlan, rocky shore, 7 
Specimens (2 ovigerous) (Apr 1980), 1 ovigerous female (Oct 1980); Isla Pajaros, 
Bay of Mazatlan, rocky shore, 4 specimens (Apr 1980); foot of Cerro de Vigia, 


VOLUME 96, NUMBER 1 73 


Bay of Mazatlan, rocky shore, 2 specimens (Dec 1980), 2 specimens (1 ovigerous) 
(Oct 1980), 3 specimens (1 ovigerous) (Nov 1980). 

Previous Eastern Pacific records.—Southern Gulf of California (Brusca 1980); 
Isla San José, Baja California Sur (Coutiere 1909). 

Range.—Scammon’s Lagoon, west coast of Baja California throughout Gulf of 
California (Wicksten in press). 

Habitat.—In the Bay of Mazatlan, one animal was taken in June 1979 among 
the calcareous tubes of the gregarious polychaete Filograna implexa on sand at 
9.5 m. All other specimens came from the lower midlittoral zone to the sublittoral 
fringe on rocky shores, where the species usually is found under sponges attached 
to stones or in small cavities. 


Automate dolichognatha De Man, 1888 


Material.—Off coast of Sinaloa (23°10'’N, 106°28’W), Van Veen grab, 1 spec- 
imen (Apr 1981). 

Previous Eastern Pacific records.—Isla Clarion, Panama, Isla Cocos, Colom- 
bia, Ecuador, and Galapagos Islands (Wicksten 1981). 

Range.—Tropical western Atlantic, tropical Indo-West Pacific region, tropical 
eastern Pacific from Sinaloa to the Galapagos Islands (Wicksten 1981). 

Habitat.—Commonly found in rocky intertidal zones (Wicksten 1981). The 
specimen from southern Sinaloa was collected by dredging on a silty sand bottom 
at 28 m. 


Automate rugosa Coutiere, 1900 


Material.—Off Punta Piaxtla, Van Veen grab, | specimen (Apr 1981); off Ma- 
zatlan, Van Veen grab, 4 specimens (Apr 1981); entrance to Estero de Urias 
(Mazatlan Harbour), sediment sample, 1 specimen (Apr 1980). 

Previous Eastern Pacific records.—Isla Cedros and Turtle Bay, Mexico; Gulf 
of Panama (Wicksten 1981; Coutiere 1909). 

Range.—Isla Cedros, west coast of Baja California; Sinaloa, Gulf of California, 
south to the Gulf of Panama. 

Habitat.—Specimens were taken from sediment samples of muddy sand col- 
lected at 27-70 m. 


Alpheus armillatus H. Milne Edwards, 1837 


Material.—Punta Chile, Bay of Mazatlan, rocky shore, 2 specimens (Oct 1980), 
1 specimen (Nov 1980); between Punta Chile and Punta Tiburon, Bay of Maza- 
tlan, under stones, 4 specimens (1 ovigerous) (Apr 1980); Estero de Urias, Ma- 
zatlan Harbour, 1 specimen (Nov 1981). 

Previous Eastern Pacific records.—Widely distributed in the Gulf of California 
(Wicksten in press). 

Range.—Tropical and warm temperature western Atlantic (Chace 1972); Gulf 
of California. 

Habitat.—The species is common in the rocky intertidal zone of the Bay of 
Mazatlan, where it lives in burrows under stones. Small specimens have been 


74 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


found under sponges and in small crevices. The species occurs from the lower 
midlittoral zone to the sublittoral fringe. 


Alpheus floridanus Kingsley, 1878 


Material.—Bay of Mazatlan, trawl, | specimen (Mar 1979), Van Veen grab, | 
specimen (Jan 1980); Off Cerro de Vigia, Bay of Mazatlan, Van Veen grab, | 
ovigerous female (Nov 1980); off Mazatlan (23°04’N, 106°16’W) trawl, | specimen 
(Mar 1981). 

Previous Eastern Pacific records.—Bahia Gonzaga, Mexico; Isla Taboga, Pan- 
ama (Wicksten in press). 

Range.—Eastern Atlantic from Guinea to Congo, Gulf of Mexico to Brazil 
(Chace 1972); Gulf of California, Panama. 

Habitat.—The animals were collected at 20-25 m on muddy sand, often with 
a significant amount of clay. Contrary to what has been reported for the western 
Atlantic coast, none of the specimens was taken intertidally. 


Alpheus leviusculus Dana, 1852 


Material.—South of Punta Chile, Bay of Mazatlan, rocky shore, 5 specimens 
(2 ovigerous) (Feb 1980; Apr 1980); Punta Chile, rocky shore, 3 specimens (Oct 
1980). 

Previous Eastern Pacific records.—Galapagos Islands (Sivertsen 1933; Hult 
1939). 

Range.—Wake Island, Canton Island (Banner and Banner 1964); Gulf of Cal- 
ifornia to Colombia (Wicksten in press), Galapagos Islands. 

Since the publication of the report of Banner and Banner (1982) on the alpheids 
of Australia, it may be necessary to consider the specimens from Sinaloa as 
members of a new subspecies that would be distinct to both A. /. leviusculus 
(Indo-West Pacific region) and A. /. bouvieri (Atlantic region). 

Habitat.—Alpheus leviusculus was collected under stones, among sand, and in 
gravel in rocky intertidal areas. It commonly burrows in the lower midlittoral 
zone and sublittoral fringe. 


Alpheus cf. A. malabaricus Fabricius, 1775 


Material.—Estero El Verde (23°25'30"N, 106°33’30"W) bottom dredge, 1 spec- 
imen (May 1979); El Tanque Canal, Caimanero Lagoon, cast net at night, 3 
specimens (May 1979); Estero de Urias, Mazatlan Harbour, in mud, 2 specimens 
(Nov 1981). 

Previous Eastern Pacific records.—Vicinity of Mazatlan (Wicksten in press). 

Range.—The specimens from Mazatlan belong to the A. malabaricus complex 
and their relationship with other members of this complex will be treated in a 
forthcoming paper (Wicksten in press). 

Habitat.—In southern Sinaloa, the species seems to be restricted to coastal 
lagoons and mangrove channels. The specimens from Estero de Urias were col- 
lected in mud, on a small flat bank bordered by mangrove trees at a salinity of 
36%o and a temperature of 31°C. Other specimens were collected on shallow 
muddy sand bottoms in a lagoon at a salinity of 37%o. 


VOLUME 96, NUMBER 1 75 


Table 1.—Species of Carideans in Different Habitats 


Rivers, estuaries and lagoons: Offshore soft or mixed bottoms: 
Alpheus malabaricus Alpheus floridanus 
Macrobrachium spp. Automate dolichognatha 
Palaemon gracilis Automate rugosa 
Palaemonetes hiltoni Latreutes antiborealis 

Intertidal-subtidal rocky areas: Leptochela serratorbita 


Lysmata californica 
Neopontonides dentiger 
Ogyrides sp. 
Periclimenes infraspinis 
Pontonia margarita 
Processa spp. 

Thor paschalis 
Trachycaris restrictus 


Alpheus armillatus 

Alpheus leviusculus 

Alpheus schmitti 
Gnathophyllum panamense 
Synalpheus apioceros sanjosei 
Synalpheus biunguiculatus 
Synalpheus digueti 
Synalpheus nobilii 


Alpheus schmitti Chace, 1972 


Material.—Punta Chile, Bay of Mazatlan, rocky shore, 2 specimens (Feb 1980). 

Previous Eastern Pacific records.—The species is known only from this record. 

Range.—Florida Keys, Antigua Island, Grenada, Tobago (Chace 1972); Ma- 
zatlan, Sinaloa. 

Habitat.—Lower midlittoral zone, under stones. 


Family Ogyrididae 
Ogyrides sp. 


Material.—Bay of Mazatlan, 58 specimens (Aug 1980—Apr 1981). 

Previous Eastern Pacific records.—Southwestern Mexico (Wicksten in press). 

Range.—Because of the uncertainty of the specific identity, the range is un- 
certain. Other specimens of Ogyrides have been taken in southwestern Mexico 
and off Southern California, U.S.A. There may be more than one species in the 
area. 

Habitat.—In the Bay of Mazatlan, the species is common on soft bottoms. 


Discussion 


At least 29 species of carideans including a least two species of Processa and 
one or more species of Ogyrides, are known now from the coast of Sinaloa. The 
fauna contains many wide-ranging species, some of which are known only from 
a few specimens. 

Certain species are characteristic of different depths or habitats (Table 1). Those 
of the coastal estuaries and lagoons are particularly interesting, having been poor- 
ly studied elsewhere in western Mexico. 

The caridean fauna of Mazatlan differs somewhat from that found in other parts 
of the Gulf of California. The small shrimp Palaemon ritteri, common at Guaymas 
and Puerto Penasco, has not been taken at Mazatlan. Thor paschalis and Hip- 
polyte williamsi, common elsewhere among Sargassum spp., also are uncommon 
or not reported from the area. Perhaps the degree of exposure to waves or the 


76 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


temperature of the water may be responsible for the difference in the caridean 
faunas. 


Literature Cited 


Abele, L. G. 1975. The macruran decapod Crustacea of Malpelo Island.—Smithsonian Contribu- 
tions to Zoology 176:69-85. 

1976. Comparative species composition and relative abundance of decapod crustaceans in 
marine habitats of Panama.—Marine Biology 38:263—278. 
, and W. K. Patton. 1976. The size of coral heads and the community biology of associated 
decapod crustaceans.—Journal of Biogeography 3:35—47. 
Banner, A. H. 1953. The Crangonidae, or snapping shrimp of Hawaii.—Pacific Science 12(1):3— 
144. 
,and D. M. Banner. 1964. Contributions to the knowledge of the aipheid shrimp of the Pacific 
Ocean, [X. Collections from the Phoenix and Line Islands.—Pacific Science 18(1):83—100. 


, and 1966. The alpheid shrimp of Thailand.—Siam Society Monographs Series, No. 
3. 168 pp. 
, and 1982. The alpheid shrimp of Australia. Part III. The remaining alpheids, prin- 


cipally the genus Alpheus and the family Ogyridae.—Records of the Australian Museum, 34(1— 
2):1-357. 

Bate, C. S. 1868. Ona new genus, with four new species, of freshwater prawns.—Proceedings of 
the Zoological Society of London, 1868:363—368. 

—. 1888. Report on the Crustacea Macrura collected by the H.M.S. Challenger during the 
years 1873-76.—Reports of the Voyages of the H.M.S. Challenger, Zoology 24:i—xc, 1—942. 

Bouvier, E. L. 1895. Sur les Palémons recueillis dans les eaux douces de la Basse-Californie par 
M. Diguet.—Bulletin du Muséum d’ Histoire Naturelle de Paris 1:159—-162. 

Bruce, A. J. 1976. Shrimps from Kenya.—Zoologische Verhandelingen, Leiden 145:1—72. 

Brusca, R. C. 1980. A handbook to the common intertidal invertebrates of the Gulf of California.— 
University of Arizona Press, Tucson, 513 pp. 

Chace, F. A., Jr. 1937. The Templeton Crocker Expedition. VII. Caridean decapod Crustacea from 
the Gulf of California and the west coast of Lower California.—Zoologica 22(2):109-138. 

——. 1962. The non-brachyuran decapod crustaceans of Clipperton Island.—Proceedings of the 
United States National Museum 113(3466):605—635. 

———. 1972. The shrimps of the Smithsonian-Bredin Caribbean Expeditions with a summary of 
the West Indian shallow-water species (Crustacea: Decapoda: Natantia).—Smithsonian Con- 
tributions to Zoology 98:1—179. 

Coutiére, H. 1900. Sur quelques Alpheidae des Cétes américaines (Collection de U.S. National 
Museum, Washington).—Comptes Rendus Hebdomadaires des Séances de l1’Académie des 
Sciences 131:356—358. 

——. 1909. The American species of snapping shrimps of the genus Synalpheus.—Proceedings 
of the United States National Museum 36(1659): 1—93. 

Dana, J. D. 1852. Crustacea. Part I. U.S. Exploring Expedition, during the years 1838, 1839, 1840, 
1841, 1842, under the command of Charles Wilkes, U.S.N., 13:1—1620. 

Fabricius, J. C. 1775. Systema Entomologiae, sistens insectorum classes, ordines, genera, species, 
adiectis synonymis, locis, descriptionibus, observationibus. Flensburg and Leipzig, 832 pp. 

Faxon, W. 1893. Reports on the dredging operations off the west coast of Central America to the 
Galapagos, to the west coast of Mexico, and in the Gulf of California, in charge of Alexander 
Agassiz, carried on by the U.S. Fish Commission steamer ‘‘Albatross”’ during 1891, Lieut. 
Commander Z. L. Tanner, U.S.N., commanding. VI. Preliminary descriptions of new species 
of Crustacea.—Bulletin of the Museum of Comparative Zoology Harvard University 24(7):149— 
220. 

Heller, C. 1862. Beitrage zur Crustaceen-Fauna des Roten Meeres. Zweiter Teil.—Sitzungsberichte 
der Akademie der Wissenschaften in Wien 44(1):241—295. 

Hendrickx, M. E., and A. M. van der Heiden. (in press). Study of the invertebrate fauna of southern 
Sinaloa, México.—In A. Ayala-Castanares, F. B. Phleger, R. Schwartzlose y A. Laguarda, 


VOLUME 96, NUMBER 1 77 


eds., The Gulf of California: Origin, Evolution, Waters, Marine Life & Natural Resources. 
Universidad Nacional Autonoma de México, Mexico City. 

Holthuis, L. B. 1950. Preliminary descriptions of twelve new species of palaemonid prawns from 

American waters (Crustacea Decapoda).—Proceedings of the Koninklijke Nederlandse Aka- 

demie van Wetenschappen 53:93-99. 

1951. A general revision of the Palaemonidae (Crustacea Decapoda Natantia) of the Amer- 
icas. I. The subfamilies Euryrhynchinae and Pontoniinae.—Occasional Papers of the Allan 
Hancock Foundation 11:1—332. 

1952a. The Palaemonidae collected by the Siboga and Snellius expeditions, with remarks 
on other species. II. Subfamily Pontoniinae.—Siboga Expeditie 39a!°: 1-249. 

1952b. A general revision of the Palaemonidae. II. The subfamily Palaemoninae.—Occa- 
sional Papers of the Allan Hancock Foundation 12:1—396. 

. 1952c. Reports of the Lund University Chile Expedition 1948-49. The Crustacea Decapoda 

Macrura of Chile.—Lunds Universitets Arsskrift, N.F. Avd. 2, 7(10):1-109. 

Hult, J. 1939. Crustacea Decapoda from the Galapagos Islands collected by Mr. Rolf Blomberg.— 
Arkiv for Zoologi 30A(5): 1-18. 

Kingsley, J. S. 1878. A synopsis of the North American species of the genus Alpheus.—Bulletin 
of the United States Geological and Geographical Survey of the Territories, United States 
Department of the Interior 4:189-199. 

Man, J. G. 1888. Bericht tiber die von Herrn Dr. J. Brock im indischen Archipel gessammelten 
Decapoden und Stomatopoden.—Archiv ftir Naturgeschichte 53(1):215-600. 

Méndez, M. 1981. Claves de identificacion y distribucion de los langostinos y camarones (Crustacea: 
Decapoda) del mar y rios de la costa del Peru.—Boletin del Instituto del Mar de Peru-Callao. 
5:1-170. 

Milne-Edwards, A. 1878. Description de quelques especes nouvelles de crustacés provenant du 
voyage aux iles du Cap-Vert de MM. Bouvier et de Cessac.—Bulletin de la Société philom- 
athématique de Paris, (7), 2:225—232. 

Milne-Edwards, H. 1837. Histoire naturelle des crustacés, comprenant l’anatomie, la physiologie 
et la classification de ces animaux. 2. Paris: Roret. 532 pp. 

Parker, R. H. 1963. Zoogeography and ecology of macroinvertebrates, particularly mollusks, in the 
Gulf of California and the continental slope off México.—Videnskabelige Meddelelse fra Dansk 
Natur-historisk Forening 126:1-178. 

Rathbun, M. J. 1902. Description of new decapod crustaceans from the west coast of North Amer- 
ica.—Proceedings of the United States National Museum 24(1272):885—905. 

Schmitt, W. L. 1921. The marine decapod Crustacea of California.—University of California, Pub- 

lications in Zoology 23:1—470. 

1939. Decapod and other Crustacea collected on the Presidential Cruise of 1938.—Smith- 
sonian Miscellaneous Collections 98(6):1—29. 

Sivertsen, E. 1933. The Norwegian Zoological Expedition to the Galapagos Islands 1925, conducted 
by Alf Wollebaek. VII. Littoral Crustacea Decapoda from the Galapagos Islands.—Meddelelser 
fra det Zoologiske Museum, Oslo 38:1—23. 

Smith, S. I. 1869. Jn Verrill, A. E. On the parasitic habits of Crustacea.—American Naturalist 3: 
239-250. 

—. 1871. List of the Crustacea collected by J. A. McNiel in Central America.—Reports of the 
Peabody Academy of Science, 1869:87-98. 

Standing, J.D. 1981. Occurrences of shrimps (Natantia: Penaeidea and Caridea) in central California 
and Oregon.—Proceedings of the Biological Society of Washington 94(3):774—786. 

Stimpson, W. 1860. Prodromus descriptionis animalium evertebratorum, quae in Expeditione ad 
Oceanum Pacificum Septentrionalem, a Republica Federata missa, Cadwaladaro Ringgold et 
Johanne Rodgers Ducibus, observavit et descripsit W. Stimpson. Pars VIII. Crustacea Ma- 
crura.—Proceedings of the Academy of Natural Sciences of Philadelphia 12:22-47. 

——. 1866. Descriptions of new genera and species of macrurous Crustacea from the west coast 
of North America.—Proceedings of the Chicago Academy of Sciences 1:46-48. 

van der Heiden, A. M., and M. E. Hendrickx. 1982. Inventario de la fauna marina y costera del 
sur de Sinaloa, México. Segundo Informe de Avance, Revisado y aumentado.—Universidad 
Nacional Autonoma de México, Instituto de Ciencias del Mar y Limnologia, Estacion Maza- 
tlan. 135 pp. 


78 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Wicksten, M. K. 1981. The species of Automate (Caridea: Alpheidae) in the eastern Pacific Ocean.— 
Proceedings of the Biological Society of Washington 94(4):1104—1109. 


—. (in press). Shallow water caridean shrimps of the Gulf of California —Occasional Papers 
of the Allan Hancock Foundation. New Series. 


, and M. Méndez. (in press). Neuvos registros de camarones carideos del Peru. 


(MEH and AMVDH) Instituto de Ciencias del Mar y Limnologia, UNAM, 
Estacion Mazatlan, P.O. Box 811, Mazatlan, Sinaloa, México; (MKW) Depart- 
ment of Biology, Texas A&M University, College Station, Texas 77843-3258. 


PROC. BIOL. SOC. WASH. 
96(1), 1983, pp. 79-83 


THREE NEW SPECIES OF OCHROTRICHIA 
(METRICHTIA) FROM CHIAPAS, MEXICO 
(TRICHOPTERA: HYDROPTILIDAE) 


Joaquin Bueno-Soria 


Abstract.—Three new species of Ochrotrichia (Metrichia) (Trichoptera: Hy- 
droptilidae), are described and figured: Ochrotrichia (M.) lacuna Ochrotrichia 
(M.) riva and Ochrotrichia (M.) avon. All species were discovered to occur 
around a waterfall named ‘“‘Cascada de Misolha’’ in Chiapas, Mexico. 


In a recent collecting trip in Mexico, I found several undescribed species of 
Ochrotrichia (M.) from a waterfall named **‘Cascada de Misolha,’’ located at 20 
km southeast from Palenque, Chiapas, on the road to Ocosingo. 

Banks (1907), described Orthotrichia nigritta from Texas, which was made the 
type of the genus Metrichia by Ross (1938). The genus was reduced to the status 
of subgenus by Flint (1968), and been revised by Flint (1972), Denning and Blickle 
(1972), and Marshall (1979). Heretofore only five species of Ochrotrichia (Me- 
trichia) have been known from Mexico: Ochrotrichia (M.) aberrans Flint, Och- 
rotrichia (M.) nigritta (Banks), Ochrtrichia (M.) quadrata Flint, Ochrotrichia 
(M.) trigonella Flint, and Ochrotrichia (M.) trispinosa Bueno. In this paper three 
new species are described. 


Ochrotrichia (M.) lacuna, new species 
Figs. 1-3 


This species appears to be related to Ochrotrichia (M.) biungulata Flint, par- 
ticularly in the shape of the claspers, but differs rather strongly in the structure 
of the aedeagus. The two, unequal, subapical hooks and internal tubule and lateral 
Spine arising just basad of them are distinctive. 

Adult.—Length of forewing, 2 mm. Color uniformly fuscous. Male abdomen 
with 2 pairs of internal sacs: | long, annulate pair between segments 5 and 6, and 
a small round pair between 6 and 7. Male genitalia: Ninth segment more than one 
and one-half times as long as high in lateral aspect, posterior margin slightly 
angulate. Cercus elongate. Dorsolateral hooks slightly decurved, basolateral scale 
very small. Claspers slightly longer than high in lateral aspect, apex slightly con- 
cave, with several small, black, peglike setae in a line on ventromesal margin. 
Aedeagus with two hooks arising subapically, one much longer than other, with 
an internal tubule and lateral process arising just basad of hooks. 

Material.—Holotype, male. MEXICO: Chiapas, “‘Cascada de Misolha,’’ 20 km 
SE from Palenque, 18 May 1981, J. Bueno and H. Velasco. Deposited in Instituto 
de Biologia UNAM (IBUNAM). 


Ochrotrichia (M.) riva, new species 
Figs. 4—6 


Although abundantly distinct, this species is related to Ochrotrichia (M.) quad- 
rata Flint by the shape in lateral view of the clasper and the hooks on the ae- 


80 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Figs. 1-3. Ochrotrichia (M.) lacuna, new species. 1, Male genitalia lateral. 2, Tenth terga and 
claspers in dorsal view. 3, Aedeagus in dorsal view. 


deagus. However, the small rectangular claspers, with a few spines at the apex, 
and the long process of the aedeagus are distinctive. 

Adult.—Length of forewing, 3 mm. Color dark-brown in alcohol. Male abdo- 
men without any sacs between the segments. Male genitalia: Ninth segment al- 
most twice as long as high in lateral aspect, posterior margin nearly vertical. 
Cercus rounded. Dorsolateral hooks long, wide, somewhat curved, basolateral 


VOLUME 96, NUMBER 1 


Figs. 4-6. Ochrotrichia (M.) riva, new species. 4, Male genitalia lateral. 5, Tenth tergum and 
claspers in dorsal view. 6, Aedeagus in dorsal view. 


scale slightly curved. Clasper rectangular in lateral aspect, posterior margin trun- 
cate and bearing a few long spines. Aedeagus with two hooks arising subapically, 
one short and very dark, other much longer and paler, with a long lateral process 
arising basad of hooks. 

Material.—Holotype, male. MEXICO: Chiapas, ‘‘Cascada de Misolha,’’ 20 km 
SE from Palenque, 18 May 1981, J. Bueno and H. Velasco (IBUNAM). Para- 
types, same data as holotype, 17 males (IBUNAM); same, but C. M. & O. S. 
Flint Jr., 2 males. Deposited in the United States National Museum (USNM). 


81 


82 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Figs. 7-9. Ochrotrichia (M.) avon, new species. 7, Male genitalia lateral. 8, Tenth tergum and 
claspers in dorsal view. 9, Aedeagus in dorsal view. 


Ochrotrichia (M.) avon, new species 
Figs. 7-9 


This species appears to be quite different from the two preceding species, 
differing rather strongly in the shape of the aedeagus. The paired slightly sepa- 


VOLUME 96, NUMBER | 83 


rated, subapical hooks, and the long internal tubule arising from the base of the 
aedeagus, are distinctive. 

Adult.—Length of forewing 2 mm. Color uniformly fuscous. Male abdomen 
with a pair of short, dark, sacs within fifth segment, opening between fifth and 
sixth. Male genitalia: Ninth segment more than one and one-half times as long 
as high in lateral aspect, posterior margin angulate. Cercus narrow and elongate. 
Dorsolateral hook distintly decurved, basolateral scale very small. Claspers clear- 
ly longer than high in lateral aspect, apex truncate, with several long setae and 
black spines on ventromesal margin. Aedeagus with two stout, black, subapical 
hooks arising almost contiguosly, one much longer than the other, with a very 
long internal tubule. 

Material.—Holotype, male. MEXICO: Chiapas, ‘‘Cascada de Misolha’’. 20 km 
SE from Palenque, 18 May 1981, J. Bueno and H. Velasco (IBUNAM). Para- 
types, same data as holotype, but C. M. & O. S. Flint Jr., | male (USNM). 


Acknowledgments 


I acknowledge the help of Dr. Oliver S. Flint, Jr. for his aid in the revision of 
the manuscript, and Dr. P. J. Spangler for the facilities used in the elaboration 
of the manuscript. 


Literature Cited 


Banks, N. 1907. New Trichoptera and Psocidae.—Journal of the New York Entomological Society 
15:162—166. 

Denning, D. G., and Blickle, R. L. 1972. A review of the genus Ochrotrichia (Trichoptera: Hy- 
droptilidae).—Annals of the Entomological Society of America 65:141—151. 

Flint, O. S., Jr. 1968. The Caddisflies of Jamaica (Trichoptera).—Bulletin of the Institute of Jamaica, 
Science Series 19:1—68. 

——. 1972. Studies of Neotropical Caddis Flies, XIII: The Genus Ochrotrichia from Mexico and 
Central America (Trichoptera: Hydroptilidae).—Smithsonian Contributions to Zoology 118: 
1-28. 

Marshall, J. E. 1979. A Revision of the genera of the Hydroptilidae (Trichoptera).—Bulletin of the 
British Museum (Natural History), Entomological Series 39(3):1—239. 

Ross, H. H. 1938. Lectotypes of North American Caddis Flies in the Museum of Comparative 
Zoology.—Psyche 45:1-61. 


Instituto de Biologia, U.N.A.M., Apartado Postal 70-153, Mexico 04510, D.F. 


PROC. BIOL. SOC. WASH. 
96(1), 1983, pp. 84-88 


STREPTOSPINIGERA HETEROSETA, A NEW GENUS AND 
SPECIES OF EUSYLLINAE (POLYCHAETA: SYLLIDAE) 
FROM THE WESTERN SHELF OF FLORIDA 


Jerry D. Kudenov 


Abstract.—Streptospinigera heteroseta is a new genus and species of Eusyl- 
linae from the western continental shelf of Florida, in the Gulf of Mexico. AI- 
though it is most closely allied to Streptosyllis in having enlarged anterior setae, 
it also exhibits similarities to both Astreptosyllis and Syllides. 


Specimens initially identified as Streptosyllis Webster and Benedict, 1884 (Po- 
lychaeta: Syllidae: Eusyllinae), were examined as part of a revision of this genus 
(Kudenov and Dorsey in preparation), and found to represent an undescribed 
genus and species. Both genera possess enlarged anterior setae, but, the new 
genus differs from Streptosyllis in having composite spinigers and capillariform 
dorsal superior simple setae. This new taxon is described herein and compared 
to Streptosyllis and the other related genera Astreptosyllis Kudenov and Dorsey, 
1982, and Syllides Orsted, 1845. 

Specimens were collected as part of the Bureau of Land Management’s Outer 
Continental Self Baseline Environmental Survey in the Gulf of Mexico (Dames 
and Moore 1979), involving Mississippi, Alabama, and Florida (MAFLA study). 
Type-specimens are deposited in the National Museum of Natural History, Smith- 
sonian Institution, Washington, D.C. 


Streptospinigera, new genus 


Type-species.—Streptospinigera heteroseta new species, by original designa- 
tion. 

Diagnosis.—Prostomium with 3 pairs of eyes, 3 antennae; palps fused basally, 
directed anteriorly and ventrally (visible from above), distally rounded, blunt. 
Pharynx unarmed. Proventriculus occupying up to 4 segments. Peristomium with 
2 pairs of tentacular cirri, these cylindrical, smooth, slightly club-shaped. Dorsal 
cirri smooth, long and cylindrical to short and subulate anteriorly, abruptly be- 
coming strongly annulate posteriorly. Ventral cirri not extending beyond para- 
podial lobes. Parapodia uniramous, supported by a single neuroaciculum, distally 
beak-shaped to truncate, anterior ones twice as thick as the rest. Notoacicula 
slender, spindle-shaped when present. Superior dorsal simple seta present, lack- 
ing distal hood, distally falcate, thick anteriorly, abruptly becoming slender, cap- 
illariform thereafter. Composite falcigers lacking distal hoods, thick anteriorly, 
abruptly becoming slender thereafter. Composite spinigers present medially and 
posteriorly, having slender shafts and prolonged blades. Inferior ventral simple 
seta absent. Pygidium terminal, with | midventral and 2 dorsolateral anal cirri. 

Remarks.—Streptospinigera is clearly most closely allied to Streptosyllis in 
having enlarged acicula accompanied by enlarged superior dorsal simple setae 
and composite falcigers in anterior-most setigers. Streptospinigera differs from 


VOLUME 96, NUMBER 1 85 


Ot mm Sb aie 0.03 mm 


Fig. 1. Streptospinigera heteroseta: a, b, d, e Holotype: a, Anterior end, dorsal view, pharynx 
partly extended; b, Right parapodium from setiger 3, dorsal view; c, Paratype: Left parapodium from 
setiger 3, view of anterior facing surface; d, Right parapodium from setiger 18, view of anterior facing 
surface; e, Pygidium, dorsal view. 


Streptosyllis in having both composite spinigers and capillariform dorsal simple 
setae, and in lacking prolonged ventral cirri and distal hoods on all setae. Strep- 
tospinigera is also somewhat similar to Astreptosyllis in that both have enlarged 
composite falcigers in anterior setigers. Astreptosyllis differs from Streptospini- 
gera in lacking both enlarged acicula and dorsal simple setae, in lacking composite 
spinigers, and in having prolonged ventral cirri. Streptospinigera is less similar 
to Syllides in that the latter lacks enlarged setae and composite spinigers. How- 
ever, Syllides and Streptospinigera have both falcate and capillariform dorsal 
simple setae, and lack prolonged ventral cirri and setal hoods. Only the type- 
species, described below, is known. 

Etymology.—The generic name derives from the Greek, streptos, meaning 
twisted, the Latin, spinatus, meaning with spines, and the Latin, gero, meaning 
to bear. Gender: feminine. 


Streptospinigera heteroseta, new species 
Figs. 1, 2 


Material examined.—FLORIDA, GULF OF MEXICO: off Tampa, MAFLA 
2207E, 27°57'00.4’N, 83°09'00.3”W, fine to very fine sand, 19 m, coll. Dames and 
Moore for BLM, Aug 1977; holotype (USNM 74489). Station 151, Florida Middle 
Ground-II, dredge, 7 Nov 1978; paratype (USNM 74490). 

Description.—A small species, measuring at least | mm long, 0.05 mm wide 
without parapodia for 23 setigers (holotype). Paratype incomplete, measuring | 
mm long, 0.08 mm wide without parapodia for 14 setigers. Body generally lacking 


86 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


5 
=e 
fe) 
ans 
0.01 mm 


Fig. 2. Steptospinigera heteroseta: a—g, j-y Holotype; h-i, Paratype: a, Aciculum, setiger 1, dorsal 
view; b, Same, setiger 2, dorsal view; c, Same, setiger 3, dorsal view; d, Same, setiger 4, dorsal 
view; e, Same, setiger 5, dorsal view; f, Same, setiger 6, dorsal view; g, Same, setiger 17, dorsal 
view; h, Notaciculum, setiger 7, dorsal view; i, Dorsal superior simple seta, setiger 3, ventrolateral 
view; Jj, Same, setiger 3, lateral view; k, Same, setiger 3, ventral view; 1, Capillariform dorsal superior 
simple seta, setiger 7, lateral view; m, Enlarged long bladed composite falciger, setiger 3, lateral view 
of anterior facing surface; n, Same, dorsolateral view of posterior facing surface; 0, Enlarged short 
bladed composite falciger, setiger 3, lateral view of anterior surface; p, Same, setiger 3; q, Same, 
setiger 4, dorsolateral view of anterior surface; r, Same, setiger 4, dorsolateral view of posterior 
surface; s, Shaft tip of enlarged, short bladed falciger, setiger 3, dorsal view; t, Slender, long bladed 
falciger, setiger 9, lateral view of anterior surface; u, Same, setiger 9, dorsal view; v, Same, setiger 
9, lateral view of anterior surface; w, Shaft tip of slender composite falciger, lateral view of anterior 
surface; x, Same, ventrolateral view of anterior surface; y, Composite spiniger, setiger 9, lateral view, 
posterior surface. 


pigmentation in alcohol; articles of annulated dorsal cirri each with dense brown 
pigment granules; with ciliary patches on prostomium, dorsal ceratophores and 
pygidium, and on raised, paired epaulettes on peristomium (Fig. la). 
Prostomium wider than long, with 3 pairs of eyes (right posterior eye as 2 
separate spots in holotype), all in hexagonal arrangement (Fig. la). Antennae 
smooth, slightly club-shaped distally; lateral antennae inserted between anterior 
pigment-patches (Fig. la); median antenna long, inserted between last 2 pairs of 
eyes (Fig. la). Palps large, directed anteriorly and ventrally, laterally incised, 
lacking palpal cirri. Pharynx partly extended in holotype, unarmed, extending 
through setiger 2 (Fig. la); number, shape of terminal papillae unknown. Pro- 


VOLUME 96, NUMBER | 87 


ventriculus barrel-shaped, with around 30 rows of muscle points, extending from 
setiger 3 through 6 (Fig. la). Peristomium a complete ring dorsally and laterally, 
with 2 pairs of tentacular cirri, these smooth, slightly club-shaped distally (Fig. 
la); with paired, mound-shaped epaulettes, these ciliated, located above each 
dorsal tentacular cirrus (Fig. Ia). 

Anterior most parapodia distally truncate (Fig. 1b, c), becoming elongate, con- 
ical posteriorly (Fig. Id). Dorsal cirri of setiger 1 long, resembling antennae and 
tentacular cirri (Fig. la); those of setigers 2—5 short, smooth, subulate to cylin- 
drical, extending beyond parapodial lobes but not setal fascicles (Fig. la); those 
from setiger 6 to end of body each with 4 articles, alternating in length as follows: 
those of setigers 6, 9, 11, 13, 16, 18 all long and well developed (Fig. la, d) while 
all others from setiger 7 short (Fig. la). Ventral cirri smooth throughout, being 
inflated in setigers 1—3 (Fig. Ic), subulate thereafter (Fig. 1d). 

Neuroacicula of setigers 2—5 (or 6 in paratype) thick-shafted, distally truncate 
(Fig. 2b-e); those from setigers |, 6-7 to end of body slender, distally beak- 
shaped (Fig. 2a, f, g). Notoaciculum accompanied by tufts of smooth capillaries 
(Fig. 2h) present from setiger 7 to end of body in paratype. Superior dorsal simple 
seta present in all setigers; those of setigers 1—5 enlarged, distally conical with a 
ventral subterminal notch and paired, lateral serrated ridges (Fig. 2i-k); those 
from setiger 6 to end of body slightly bent, slender setae with transversely ser- 
rated ventral cutting surfaces (Fig. 21). 

Setigers 1-5 each with 8-10 thick-shafted composite falcigers including 2-3 
having long and 6—7 having short unidentate blades (Fig. 2m-—r), all with saw- 
tooth cutting margins, lacking distal hoods and sheaths; shaft tips with dorsal 
superior branch terminating in an incised lobe (Fig. 2s) with either paired subdistal 
denticles (Fig. 2p, s) or an unpaired median flap (Fig. 2n, q, r); with ventral 
inferior branch entire, rounded (Fig. 2s); shaft tips of long bladed falcigers some- 
times forming clear, cup-shaped sockets for blade. (Fig. 2m, n) 

Setigers 6 to end of body each with 7-10 slender composite setae per fascicle 
including 2—3 superior spinigers each having long, minutely serrated blades (Fig. 
2y) and 5-7 falcigers having unidentate blades with finely serrated cutting margins 
(Fig. 2t—v), sometimes with pronounced basal spurs (Fig. 2t), all becoming shorter 
ventrally within a fascicle; shaft tips similar to those of setigers 1-5 except dorsal 
superior branch inconspicuously incised (Fig. 2w, x), generally smooth, lacking 
subterminal denticles. 

Pygidium short, equalling length of last 2 prepygidial segments, with anal cirri 
including | short midventral and 2 long dorsolateral cirri (left one missing, scar 
conspicuous); all smooth (Fig. le). 

Remarks.—Streptospinigera heteroseta differs from all known species of 
Streptosyllis in having composite spinigers and two kinds of superior dorsal sim- 
ple setae. It is most closely related to the Streptosyllis websteri-S. bidentata-—S. 
cryptopalpa species subgroup in that compound setae of anterior fascicles have 
both long and short appendages. Streptospinigera heteroseta differs from the 
species of this subgroup in having anteriorly, rather than ventrally, directed palps. 
Streptosyllis latipalpa is similar to Streptospinigera heteroseta in having similarly 
oriented palps. However, Streptosyllis latipalpa has only short-bladed composite 
setae in anterior fascicles. 


88 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Etymology.—The specific name, heteroseta, derives from the Greek, heteros, 
meaning different, and the Latin, seta, meaning bristles; it is used as a noun in 
apposition. 


Acknowledgments 
I am indebted to J. M. Uebelacker and P. G. Johnson of Barry A. Vittor and 
Associates, Mobile, Alabama, for allowing me to examine these syllids. I am also 
grateful to K. Fauchald, National Museum of Natural History, for critically com- 
menting on this paper, and to S. J. Williams, Allan Hancock Foundation, for 
providing technical assistance. 


Literature Cited 


Dames, T., and W. Moore. 1979. Final report—The Mississippi, Alabama, Florida, Outer Conti- 
nental Shelf Baseline Environmental Survey MAFLA 1977/1978, Vols. I-A,B, for the U.S. 
Bureau of Land Management, Washington, D.C. NTIS PB-294 288. New Orleans, Louisiana. 

Kudenov, J. D., and J. H. Dorsey. 1982. Astreptosyllis acrassiseta, a new genus and species of 

the subfamily Eusyllinae (Polychaeta: Syllidae) from Australia.—Proceedings of the Biological 

Society of Washington 95:575—578. 

. A revision of the genus Streptosyllis (Polychaeta: Syllidae: Eusyllinae).—Proceedings of the 

Biological Society of Washington. In preparation. 

Orsted, A. S. 1845. Fortegnelse over Dyr, samlede i Christianiafjord ved Drgbak fra 21-24 July 
1844.—Naturhistorisk Tidssrkrift K@benhavn Series 2, 1:400—427. 

Webster, H. E., and J. E. Benedict. 1884. The Annelida Chaetopoda from Provincetown and 
Wellfleet, Mass.—Annual Report of the Commissioner for Fish and Fisheries for 1881:699— 
747, 8 pls. 


Department of Biological Sciences, University of Alaska, Anchorage, 3221 
Providence Drive, Anchorage, Alaska 99508. 


PROC. BIOL. SOC. WASH. 
96(1), 1983, pp. 89-102 


THE IDENTITY OF PETROLISTHES MARGINATUS 
STIMPSON, 1859, AND THE DESCRIPTION OF 
PETROLISTHES DISSIMULATUS, N. SP. 
(CRUSTACEA: DECAPODA: PORCELLANIDAE) 


Robert H. Gore 


Abstract.—Petrolisthes marginatus, briefly described by Stimpson, but erro- 
neously illustrated as such by Benedict, is shown to be a composite of three 
nominal species, Petrolisthes marginatus s. s., Petrolisthes cessacii (A. Milne 
Edwards) a junior subjective synonym, and an undescribed species (P. margina- 
tus sensu Benedict). The latter form is described and illustrated as Petrolisthes 
dissimulatus. 


Petrolisthes marginatus Stimpson, 1859, is a member of a genus of cryptic, 
often fast-moving porcellanid crabs that inhabit crevices and interstices in reefs 
and other hard, more or less permanent substrata. Stimpson briefly described but 
did not illustrate P. marginatus, using a single specimen collected at Barbados 
by a Mr. Gill. Stimpson’s original specimen was presumably destroyed in the 
Great Chicago Fire (see Evans 1967), and apparently no syntypic material was 
deposited by Stimpson in any other museums. 

Petrolisthes marginatus was nevertheless considered to be well-characterized, 
based on a combination of Stimpson’s brief original description, plus a subsequent 
illustration asserted to be this species by Benedict (1901). The species is tropical, 
but not very common, being recorded from Fernando de Noronha (Pocock 1890), 
the Caribbean Sea (Young 1900), a few other localities in the western Atlantic 
(Haig 1956), and eventually but erroneously from several areas in the tropical 
eastern Pacific (Haig 1960). Specimens from the latter region were re-examined 
by Chace (1962) and placed in a new species, Petrolisthes haigae, which became 
the trans-Panamanian geminate form of P. marginatus sensu Stimpson (see be- 
low). 

Benedict’s (1901) illustration that he attributed to P. marginatus was that of a 
porcellanid collected at Ponce, Puerto Rico. Although his perfunctory description 
added little to Stimpson’s sparse characterization, the accompanying figure was 
more detailed, and showed a small crab with quadridentate cheliped meri, lacking 
walking legs, and bearing two small clearly visible epibranchial spines. These 
spines had not been mentioned by Stimpson in his original description, but later 
authors (e.g. Schmitt 1924a, b, 1935; Chace 1956, 1962; Gore 1974) followed 
Benedict’s lead and used them as criteria in identifying, or distinguishing between 
P. marginatus sensu Benedict, and other closely related but single-spined con- 
geners such as Petrolisthes amoenus (Guérin, 1855), Petrolisthes haigae Chace, 
1962, or Petrolisthes cessacii (A. Milne Edwards, 1878). 

Petrolisthes cessacii has long been considered to be closely related to Petro- 
listhes marginatus. The original description of the former by A. Milne Edwards, 
as Porcellana Cessacii, was also without an illustration and characterized a crab 


90 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


attaining a relatively large size (17 X 16 mm, cl X cw), possessing a strongly 
advanced frontal region produced into a point (‘‘affecte la forme d’un bec’’), a 
cheliped carpus having three or four denticulations on its anterior margin and a 
large spine terminating its posterior margin. There was no mention of any epi- 
branchial spination, and the remaining characters in the description could easily 
have been applied to any of several species of porcellanid crabs known at that 
time. For the purposes of this paper, however, the large size, advanced front, 
tri- or quadridentate cheliped carpus with a strong posterior spine provide evi- 
dence of similarity with Petrolisthes marginatus Stimpson (see below). 

Balss (1914) provided a brief figure of P. cessacii, but Chace’s (1956) illustration 
of the species was by far the most detailed. Literature records from Chace’s 
study suggested that P. cessacii was a rather common porcellanid crab, appar- 
ently restricted to the tropical-subtropical eastern Atlantic Ocean. However, Gore 
(1974) noted a range extension for the species to the western Caribbean Sea, and 
in the same study placed Pocock’s (1890) specimens of P. marginatus from Fer- 
nando de Noronha, Brazil, into synonymy with P. cessacii. Gore considered 
Fernando de Noronha as the first, albeit ‘‘misidentified,’’ western Atlantic record 
for Milne Edwards’ species. This action followed Coelho (1970) who previously 
had listed P. cessacii from the coast of Brazil without further comment. As a 
comparison of the illustrations provided by Chace and by Gore shows, the west- 
ern African and western Caribbean forms are quite similar. 

Most recently, R. B. Manning and F. A. Chace (in prep.) identified some 
specimens of porcellanid crabs collected from Ascension Island in the South 
Central Atlantic as P. marginatus. A comparison of this material with specimens 
of P. cessacii convinced them (in litt.) that Petrolisthes cessacii (A. Milne Ed- 
wards, 1878) was a junior synonym of Petrolisthes marginatus Stimpson, 1859. 
This being so, specimens identified as P. cessacii from Brazil (Coelho 1970; 
Coelho and Araujo Ramos 1972), Colombia (Werding 1977), the western Carib- 
bean Sea (Gore 1974), and the coast of Mexico (Rickner 1975) were probably P. 
marginatus as well. What was more important, however, was that the specimen 
reported and illustrated by Benedict (1901) as P. marginatus, and perpetuated as 
such by subsequent authors, was now seen to differ substantially from Stimpson’s 
description of the species, and because it was misidentified it would require a 
name. 

Manning and Chace’s conclusions were briefly incorporated in a report on 
Central American Porcellanidae by Gore (1982), in which the first occurrence 
of P. marginatus on the Caribbean coast of Panama was noted. In the present 
report, I provide a revised synonymy for Petrolisthes marginatus Stimpson, ex- 
pand Stimpson’s original description by including Chace’s (1956) diagnosis for P. 
cessacii, and give a detailed description and illustration for the new species pre- 
viously misidentified by Benedict. 

In the taxonomic treatment that follows, carapace length (cl) precedes carapace 
width (cw) measured in mm across the longest or widest parts of the cephalotho- 
rax. Paratypic material of the new species from Colombia, part of the personal 
collection of Dr. Bernd Werding, was returned to him; the remaining paratypic 
and holotypic material has been returned to the National Museum of Natural 
History, Washington, D.C. Repository abbreviations follow the appropriate ma- 
terial and are explained in the Acknowledgment section. 


VOLUME 9, NUMBER | 9| 


Petrolisthes marginatus Stimpson, 1859 
Fig. 1 


Petrolisthes marginatus Stimpson, 1858:227 [nomen nudum]; 1859:74 [p. 28 on 
separate].—Pocock, 1890:513.—Young, 1900:394.—Haig, 1956:26 [in part, in- 
cluding color notes and reference to Buccoo Reef specimen]; 1962:176 [in part, 
1 2, “Danish West Indies’’; not Virgin Islands specimens].—Gore, 1982:17. 
Not Nobili, 1897:4 [=Petrolisthes armatus (Gibbes, 1850), fide Nobili, 1901: 
le 

Porcellana Cessacii A. Milne-Edwards, 1878:229 [p. 10 on separate]. 

Petrolisthes Cessaci.—A. Milne Edwards and Bouvier, 1900:346. 

Petrolisthes cessaci.—Balss, 1914:101, fig. 6.—Lebour, 1959:128, 136, fig. 10 
[megalopa, identification by implication].'—Gauld, 1960:64. 

Petrolisthes Cecoci.—Balss, 1914:100 [erroneous spelling, legend, fig. 6]. 

Petrolisthes cessacii.—Sourie, 1954:84, 112, 236, 239, 253, 256, 294, 295, 304.— 
Chace, 1956:14, fig. 4A—E.—Holthuis and Manning, 1970:242 [discussion], 243.— 
Coelho, 1970:233 [listed].—Coelho and Araujo Ramos, 1972:173.—Gore, 1974: 
710, fig. 4; 1982:17 [discussion].—Rickner, 1975:163.—Werding, 1977:176, 197, 
199, fig. 19. 

Petrolisthes armatus.—Miers, 1881:432.—Balss, 1922:108 [listed].—Haig, 1962: 
178 [discussion].—Coelho, 1966:55 [references to Pocock and Nobili].—Faus- 
to-Filho, 1974:8. Not Petrolisthes armatus (Gibbes, 1850). 


Material.—MEXICO: Vera Cruz, Isla de Lobos; coral reef; 8 June 1973; coll. 
J. A. Rickner 1 6; AHF 1974-9.—PANAMA (Atlantic): Golfo de San Blas, Pico 
Feo; Thalassia flat; 0-1 m; 7 Apr 1973; coll. Newman and party; | 6, 1 2 
ovigerous; USNM 292580.—OLD PROVIDENCE ISLAND; 13°31.55'N, 
81°20.55'W; 0-1 m; 30 Jan 1972; R/V John Elliott Pillsbury, sta P-1350; 1 6,1 2 
ovigerous; IRCZM 89:3741.—Same station; 3 6¢, 1 2; UMML 32:5515.—Same 
Station; 3 2 2 ovigerous; RMNH 30407.—Same locality; low cay; boulders; 0-1 
m; 14 Dec 1980; coll. B. Werding; | juvenile; personal collection COLOMBIA: 
Santa Marta, Aguja; Playa Brava; under stones; 1.5 m; Dec 1978; coll. B. Werd- 
ing; 2 6d, 2 22 ovigerous; personal collection.—Same area; Morro Grande; 
intertidal, under stones; May 1980; coll. B. Werding; 3 66,9 2 2 (7 ovigerous); 
personal collection.—Same area; Burrucuca; intertidal, among stones; 12 Jul 1976; 
coll. B. Werding; | ¢, 1 2; personal collection.—Same locality; shore; Nov 1976; 
coll. B. Werding; | 6; AHF 726-01.—Same area; Punta de Betin; intertidal, 
among stones; Nov 1976; B. coll. Werding; | 6; personal collection.—Same area; 
Tagange; intertidal; Feb 1976; coll. B. Werding; | 6; personal collection.—Same 
area; Islas Rosario; May 1977; coll. B. Werding; | 6; personal collection.— 
CURACAO: southwest coast, west of Willemstad; under stones on shore; Jan— 
Feb 1957; coll. B. van Bergeyk; 1 2 ovigerous; LBH 1151.—Same; Fuik Bay; 


1 The megalopa Lebour attributed to Petrolisthes cessacii may not belong to this species, because 
the figure she provided shows no epibranchial spines. Megalopae of other Petrolisthes species that I 
have examined invariably exhibit such spines if they occur in the species in the adult stage. The 
megalopa illustrated by Lebour could therefore be one of several species of Petrolisthes which occur 
on, or in the vicinity of, the west African coast. On the possibility that Lebour overlooked the 
postlarval epibranchial spines, and without evidence attributing this megalopa to another species, I 
will provisionally maintain Lebour’s identification within the synonymy of P. marginatus. 


92 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


ee oe 


Fig. 1. Petrolisthes marginatus. A, Male, R/V John Elliot Pillsbury, Sta. P-1350, Old Providence 
Island, UMML 32:5515; B, Epibranchial and lateral margin of carapace, showing spination; C, Right 
cheliped, posteroventral view of marginal spines; D, Left cheliped, mesioventral view, note single 
meral spine. Scale A = 5 mm; B—D = 3 mm. 


NW landing; intertidal, among stones; 0-1 m; 13 Jan 1957; coll. L. B. Holthuis; 
1 2 crushed; LBH Nr 1051.—KLEIN BONAIRE: Old landing by Bonaire; coral 
debris on shore; 0-1 m; 17 Oct and 8 Nov 1930; coll. P. W. Hummelinck; | 6, 
1 2 crushed; PWH 1049a.—Same locality; east coast by landing place; rock, 
sand, tidal zone; 13 Sep 1948; coll. P. W. Hummelinck; 2 66, 1 9 ovigerous, 1 
juvenile; PWH 1049a.—*‘DANISH WEST INDIES”: 15 Dec 1911; col.? C. Meng; 
1 2; no other data; UZMC collection. — VIRGIN ISLANDS: St. John, Europa 
Bay; Feb 1959; coll. Randall and Kumpf; 1 6, 1 2 ovigerous; UMML 32:1595.— 
SABA, NETHERLANDS ANTILLES: E. Fort Bay; rocks, tidal zone and lower; 
21 Jul 1949; coll. P. W. Hummelinck; 1 2 ovigerous; RMNH 8458.—AVES (Islote 
Aves): W of Dominica; N reef, rocks, tidal zone; 12 May 1949; coll. P. W. 
Hummelinck; 1 2; RMNH 8456.—TRINIDAD: Maracas Bay; southwest corner; 
intertidal; 18 Jul 1968; R/V John Elliott Pillsbury, sta P-701; 1 2 ovigerous; 
UMML 32:5481.—TOBAGO (British West Indies): Buccoo Reef; shore; 20 Apr 
1939; Velero III, Hancock Expedition; 1 ¢; A41-39.—BRAZIL: Fernando 
(de) Noronha; coll. H. M. Ridley; no other data; 3 66; BMNH 1888.19.—WEST 
AFRICA: Senegal; Pointe des Almadies; 30 Jul 1973; 1 6, 1 2 ovigerous; MNHN 


VOLUME 96, NUMBER 1 93 


Ga 551.—Same; Gulf of Guinea; Fernando Poo; 3°45’N, 8°48’E; R/V John Elliott 
Pillsbury, sta P-258 shore; 15 May 1965; 2 646, 3 22; RMNH 23976.—Same; 
Annobon; 1°25’S, 5°38’E; R/V John Elliott Pillsbury, sta 271, shore; 19 May 1965; 
736,3 2 (2 ovigerous); RMNH 23975. 

Diagnosis.—Carapace about as wide as long, covered with short pubescence, 
usually distinct, occasionally almost invisible, smooth except for few faint rugae 
posterolaterally; single sharp or blunt epibranchial spine, rarely with nub of sec- 
ond immediately behind. Front with prominent median lobe, projecting well be- 
yond lateral lobes, latter nearly transverse, orbital margins oblique. Merus of 
chelipeds armed anteroventrally with single, usually acute spine; carpus with 3- 
5, usually 4, minutely serrate, spine-tipped, nearly equal-sized teeth; posterior 
margin with 2-6 oblique rugae produced apically into spines, strong, hooked, 
bifid spine at posterodistal angle. Chelae pubescent, bearing distinct granular 
ridge from dactylar to chelar bases, separating flattened upper part of manus from 
slanting outer part. Walking legs spinulous on anterior margins of meri, numbers 
ranging from 4-8, posterodistal angles produced into | or 2 teeth on legs 1 and 
2, unarmed on walking leg 3. (Modified from Chace 1956:15.) 

Distribution.—Western Africa from Cape Verde Islands to Annobon, on the 
mainland from Senegal to Ghana; Ascension Island, South Central Atlantic Ocean; 
Fernando de Noronha and Sao Luiz, Brazil, north to Trinidad, Tobago Island, 
vicinity of Santa Marta, Colombia, Curacao, Bonaire, Old Providence Island, 
Panama, Mexico, and the northern Leeward Islands. In the littoral and shallow 
sublittoral to about 2 m. 

Remarks.—With the placing of Petrolisthes cessacii into synonymy, Petro- 
listhes marginatus becomes another of a small group of essentially tropical por- 
cellanid crabs in the genus Petrolisthes having amphi-oceanic or amphi-continen- 
tal distribution. In addition to P. marginatus, now known from western Africa, 
Ascension Island, eastern South America and the Caribbean Sea, the group in- 
cludes Petrolisthes armatus (Gibbes, 1850) from western Africa, eastern tropical 
North, Central and South America, and the tropical eastern Pacific; Petrolisthes 
galathinus (Bosc, 1802) from warm-temperate and tropical eastern North Amer- 
ica, tropical Central and South America and the eastern Pacific; Petrolisthes 
tonsorius Haig, 1960 from the southwestern Caribbean Sea and tropical eastern 
Pacific; and Petrolisthes tridentatus Stimpson, 1859, from the Caribbean Sea, and 
tropical eastern South America and eastern Pacific. These species have presum- 
ably been able to cross oceanic or isthmian barriers either by dispersal or by 
vicariance events. The geographically separated populations at least have not 
undergone speciation recognizable through presently employed morphological 
criteria. At least 14 other species of Petrolisthes have extensive tropical distri- 
butions encompassing the western Indian Ocean and to either Japan or to Samoa or 
Hawaii in the Pacific Ocean (Haig 1964, 1974, in litt.). 

The relegation of P. cessacii into junior synonymy with P. marginatus appears 
to be justified at present based on morphology. Whether the eastern and western 
Atlantic populations are reproductively isolated or will remain so, given the po- 
tential for long distance larval dispersal, remains uncertain. It would be extremely 
interesting, for example, to see if differences in larval morphology are exhibited 
among the African, Ascension Island, and western Atlantic populations. 


94 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 2. Petrolisthes dissimulatus, holotype, ovigerous female, St. John, Virgin Islands, USNM 
190893. Scale line = 3 mm. 


Petrolisthes dissimulatus, new species 
Figs. 2, 3, 4 


Petrolisthes marginatus.—Benedict, 1901:134, pl. 3, fig. 1.—Schmitt, 1924a:73; 
1924b:88; 1935:185, 187.—Boschma, 1931:374.—Chace, 1956: 24 [discussion]; 
1962:622 [discussion].—Haig, 1956:17, 26 (in part, Curacao specimen only); 
1962:176 (in part, except | 2, “‘Danish West Indies’’).—Gore, 1974:711 [dis- 
cussion]. Not P. marginatus Stimpson, 1859. 


Holotype.—1 @, ovigerous; St. John, Virgin Islands, coral reef of lagoon point, 
west side of bay; 6 Apr 1937; W. L. Schmitt; USNM 190893. 

Paratypes.—2 36, same data as holotype; USNM 190894. PUERTO RICO: 
Playa de Ponce; 1899; USFC Steamer Fish Hawk; J. E. Benedict; 1 2; USNM 
42351.—VIRGIN ISLANDS: St. Thomas; 10 Nov 1917?; no other data; UZMC 
collection. —‘‘DANISH WEST INDIES”: 15 Dec 1911; C. Meng; 3 d6, 2 29 
(1 ovigerous); no other data; UZMC collection —BARBADOS (Univ. Iowa Bar- 
bados-Antigua Exped.): 15 May 1918; W. L. Schmitt; 1 6, 2 22 ovigerous; 
USNM 57971.—Same; from coral head; 4 Jun 1918; W. L. Schmitt; 1 6; USNM 
57972.—Same; Okra Reef; 13 May 1918; W. L. Schmitt; | ¢, 2 2 2 ovigerous, 
1 cheliped; USNM 68648.—Same; 15 May 1918; W. L. Schmitt; 4 6d, 1 2 
ovigerous; USNM 68649.—Same; coral heads; 4 Jun 1918; W. L. Schmitt; 1 9; 
USNM 68650.—Same; old coral; 31 May 1918; W. L. Schmitt; 1 ¢, 2 22 ovi- 
gerous; USNM 68651.—CURACAO: Santa Marta Bay, by St. Nicolaas; coral 
blocks, open sea shore; 4 Feb 1957; coll. L. B. Holthuis; 1 6; LBH nr. 1083.— 


VOLUME 96, NUMBER 1 95 


Same; Piscadera Bay; sand and muddy sand with stones, algae, sponges, etc.; 0- 
1.5 m; 12 Nov and Dec 1956; coll. L. B. Holthuis; 1 ¢, 2 2 2 (1 ovigerous); LBH 
nr. 1002.—Same; Willemstad; strand of Marie Pompoen; among stones; 26 Dec 
1956; coll. L. B. Holthuis; 3 66; LBH nr. 1028.—Same; Fuik Bay; NW landing; 
among stones; 0-1 m; 13 Jan 1957; coll. L. B. Holthuis; 6 6d, 3 22; LBH nr. 
1051.—Same; Vista Alegre; shore; 23 Apr 1939; **Velero III’’ Hancock Expedi- 
tion; | 2 ovigerous; USNM 98105 [A46-39]—COLOMBIA: Santa Marta; 
Aguja, Playa Brava; under stones; 1.5 m; Dec 1978; coll. B. Werding; 1 5, 1 9; 
personal collection.—Same; Islas Rosario; May 1977; coll. B. Werding; 1 6, 3 
2 2 (2 ovigerous); personal collection. 

Measurements.—Holotype, 4.1 x 4.0 mm; paratype males, 1.8 x 1.8 to 6.5 x 
5.9 mm; paratype females, 3.1 x 3.0 to 5.8 x 5.4 mm; ovigerous females, 3.9 x 
3.6 to 5.7 < 5.6 mm. 

Diagnosis.—Carapace rounded, smooth, pubescent; frontal region produced, 
trilobate, truncate anteriorly, prominent rounded denticulate median lobe, lateral 
lobes rectangularly rounded, serrated; orbital margin nearly normal to plane of 
lateral lobe; 2 epibranchial spines; manus broad, flattened, distinct longitudinal 
row of low tubercles extending along outer surface from dactylar to chela bases; 
merus with 2 sharp spines on anteroventral margin; carpus less than twice as long 
as wide, anterior margin with 4 serrated teeth having curved needlelike tips, 
posterior margin distal extension a single curved spine; walking legs spinulous 
on anterior margins of meri, posterodistal angles of legs 1 and 2 produced into 
sharp tooth, that of leg 3 unarmed. 

Description.—Cephalothorax subcircular to slightly rectangular, longer than 
broad, flattened posteriorly, smooth, covered with fine pubescence predominantly 
on frontal, and anterior and posterior branchial areas; frontal region produced, 
trilobate, lateral lobes rectangularly rounded, minutely dentate especially at inner 
orbital angle, anterior margins nearly transverse, front thus appearing truncate; 
prominent, rounded, slightly deflexed, denticulate, median lobe, subequal to lat- 
eral lobes, shallow, median sulcus extending posteriorly to paired, slightly ele- 
vated protogastric lobes; orbital margin nearly straight, strongly concave poste- 
riorly, forming parabolic arc, mesial and posterior margins appearing as rounded 
right angle; no supraocular spines or spinules, but minute denticles often present; 
outer orbital angle a sharp spine directed obliquely outward, followed by small 
serrations decreasing in size posteriorly; epibranchial angle with single large curved 
acute spine above, a second, often less acute, occasionally only a nub, immedi- 
ately behind; remainder of carapace unarmed; lateral margins from epibranchial 
angle to posterior branchial region bearing thin ridge forming general dorsal out- 
line of cephalothorax. 

First movable antennal article with large, lamellar projection bearing several 
smaller teeth leading to strong single, or bifid, spine at tip, second with longitu- 
dinal row of small acute conical denticles or tubercles, third smooth, rounded; 
flagellar articles lightly setose. 

Third maxilliped ischium exhibiting several raised, granular, transverse ridges, 
these appearing more like rugae on merus; mesial triangular projection of latter 
with distinct spinule at apex; remaining articles lightly rugose or smooth. 

Cheliped merus rough, granular, large serrated, spinelike tooth on anterior 
margin, followed by pair of small, curved spinules on posterodorsal margin, sec- 


96 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 3. Petrolisthes dissimulatus, Benedict's specimen, paratype female, Playa de Ponce, Puerto 
Rico; USNM 42351. Scale line = 3 mm. 


ond pair (rarely 3), more closely spaced, often extremely sharp or needlelike on 
anteroventral margin; article pubescence sparse but produced into several long 
tufts on posterior margin; carpus about 1.4 x longer than wide, covered dorsally 
with very low, rounded granules scarcely raised from surface of article, plus 
isolated raised tubercles interspersed throughout, all often obscured by thick, fine 
pubescence; anterior margin bearing 4 strong, serrated, distinctly curved teeth 
which decrease in size distally, and having spinelike or needlelike tips; prominent 
medial longitudinal ridge formed by irregular row of large, flattened tubercles 
extends to medial distal margin, many of these tubercles carrying fanlike row of 
simple setae; posterior margin with series of 6—11 elongate, raised, ridgelike se- 
tose rugae, those proximally ending in single, thin, upright spine pointing dorsally, 
those more distally becoming wider, increasingly horizontal in orientation, that 
of distalmost extension at posterodistal angle a single, strongly curved spine; 
another smaller spine adjacent on posterodorsal medial lobe; manus broad, flat- 
tened, with low granules nearly obscured by fine pubescence as on carpus; lon- 
gitudinal crest of enlarged, flattened tubercles on upper quarter of outer surface, 
extending from dactylar to carpal junctions, second row, more elongated and 
flattened, defining proximal upper margin, both rows forming oblique isoceles 
triangle; ventral margin of hand a combination of small, conical teeth or crenulate 
tubercles in irregular double row to tip of fixed finger; another single row defines 
anterior cutting edge of latter, becoming completely smooth at finger base; gape 
interiorly with thick tuft of pubescence; movable finger trihedral, upper margin 
defined by a longitudinal row of enlarged, adpressed or imbricate teeth, coupled 
with less distinct adjacent second row, both joining that on manus; dactylar tip 
curved, crossing over interiorly to similarly curved tip of fixed finger; cutting 
edges of both without noticeable exterior gape. 


VOLUME 96, NUMBER 1 97 


BILE LG 


“ay UC, eve 
i ee Ag: 


CQ -—_—_—_——_ ae 
A,B,D-F -——_—+ G 


Fig. 4. Petrolisthes dissimulatus. A, C, D, Holotype female, USNM 190893; B, Paratype male, 
St. John’s, Virgin Islands, USNM 190894; E—G, Benedict's specimen, paratype female, Playa de 
Ponce, Puerto Rico; USNM 42351. A, E, Epibranchial and lateral margin of carapace, showing 
spination; B, F, Right cheliped, posteroventral view of marginal spines; C, G, Left cheliped, me- 
sioventral view; note double meral spines; D, Left maxilliped 3. Scale A, B, D, F = 1 mm; C, G, = 
3 mm. 


98 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Walking legs smooth to faintly rugose; anterior margin of meri with many fine 
plumose hairs, these less numerous on carpus, all segments with scattered long, 
non-plumose setae; anterior margins of meri variably spied, that of first and 
second with 5-9, third with 3-4 (rarely 7) sharp, curved spinules; posterodistal 
angle of legs 1 and 2 produced into single, occasionally double tooth, third un- 
armed; distal dorsal margin of carpi with single strong tooth or spine; propodi 
with 2 widely-spaced ventral spines, stronger pair or triad at distoventral margin; 
dactyls with 3 accessory spinules nearly equal in length to each other, subequal 
to dactylar tip; ventral bases of all pereopods with sharp spinule posteriorly, best 
seen in individuals having autotomized appendages. Telson with 7 plates. 

Variations.—The most consistent variation was in the sharpness of spines and 
spinules. On the meri and carpi of the chelipeds this ranged from distinctly curved 
and needlelike, to merely acute, to somewhat blunt. Signs of wear and breakage 
were often seen. Although the epibranchial spines were usually acute, in several 
instances the second and smaller spine was less sharp, often blunt, or appearing 
as a nub, quite difficult to discern in the surrounding pubescence. Some variation 
was also seen in the pilosity on the carapace and chelipeds but is probably at- 
tributable to wear and exigencies of long years of perservation. 

Color.—All specimens had faded in alcohol to creamy white. According to 
Benedict (1901) his specimen was pink when first examined. Schmitt (1924b) 
stated that Barbadoan specimens in alcohol were salmon pink, marked with red- 
dish flecks along the margins and tubercles of the chelipeds, and with red bands 
on the carpus and propodus of the walking legs. 

Ecology.—The species has been collected from littoral and shallow sublittoral 
coralline rubble and rocky habitats associated with coral reefs. Ovigerous females 
are known only from April and May. 

Distribution.—The Caribbean Sea; the Antilles from Puerto Rico, the Virgin 
Islands, Barbados, and Curagao; Colombia from Santa Marta and Islas Rosario. 

Relationships.—Petrolisthes dissimulatus exhibits some relationship to several 
western Atlantic or eastern Pacific porcellanids. It superficially resembles the 
Caribbean species Petrolisthes amoenus (Guérin) in general carapace shape and 
cheliped morphology and armature, but differs in lacking a supraocular spine and 
the smaller spinules on the lateral margins of the carapace seen in P. amoenus. 
Petrolisthes amoenus is also less pubescent, and does not exhibit the distinctive 
granular ridges on the upper outer quarter of the manus. The new species bears 
a close resemblance to what may be its eastern Pacific geminate species, Petrolis- 
thes hirtispinosus Lockington, 1878. Both possess granular ridges on the manus, 
but P. dissimulatus can be easily distinguished from P. hirtispinosus (and P. 
amoenus as well) by the double epibranchial spines, and the truncate, nearly 
transverse frontal margin. The new species also shows a general resemblance to 
Petrolisthes monodi Chace, 1956, a somewhat variable species from northwestern 
Africa, but can be separated using the characters previously delineated for dis- 
tinguishing it from New World relatives. 

Etymology.—The specific epithet is Latin, meaning feigned, disguised, or hid- 
den, and refers to the fact that the new species was confused with P. marginatus 
for sO many years. 

Discussion.—Five morphological characters were emphasized both in Stimp- 
son’s original description of P. marginatus, and in Benedict’s description and 


VOLUME 96, NUMBER 1 99 


illustration of what is now P. dissimulatus. These are 1) the shape and prominence 
of the frontal region, 2) the shape and size of the cheliped carpal teeth, 3) the 
presence of a ridge on the upper surface of the manus, 4) the color of the spec- 
imens, and 5) the overall size of the material at hand. 

According to Stimpson, in P. marginatus the median lobe of the front is prom- 
inent, the four teeth on the anterior margin of the cheliped carpus are equal-sized 
and very sharp, the ridge extending from the anterior angle of the carpus to the 
inner base of the finger on the chelipeds is of moderate prominence, the color of 
the species was deep purplish-crimson, and the carapace length was ‘‘about half 
an inch”’ [ca. 12 mm]. 

In Benedict’s description and illustration the outline of the front from the me- 
dian lobe to the angle of the orbit is straight, the cheliped carpus bears four sharp 
marginal teeth which decrease in size distally, a granular ridge extends from the 
gape of the fingers to the anterior edge of the carpus, the color of the specimen 
was pink, and the carapace length of the illustrated specimen is approximately 4 
mm, based on a listed magnification of 312. The female specimen of P. dissim- 
ulatus from Playa de Ponce (USNM 42351) is 3.9 x 3.6 mm and agrees with 
the figured specimen not only in general measurements, but also in carapace and 
cheliped morphology, even down to the damaged distalmost marginal tooth on 
the right cheliped, and is thus the specimen Benedict studied (see Figs. 2, 3, 4). 

Although P. dissimulatus agrees in a very general way with Stimpson’s original 
description of P. marginatus, when the two forms are placed side by side the 
differences are immediately seen (compare Figs. | and 4 herein). For example, 
in P. marginatus the median lobe of the front is always quite prominent and much 
narrower than that of P. dissimulatus. The lateral margins of the front sloee more 
or less obliquely backward to the inner orbital angle in P. marginatus whereas 
in P. dissimulatus they are nearly transverse or “‘straight’’ as noted by Benedict 
(1901) and Schmitt (1935). The cheliped carpal teeth in P. marginatus are sharply 
pointed, equal-sized or nearly so, or diminish only gradually in size. In P. dis- 
simulatus these same teeth are not only sharply pointed but may have needlelike 
tips which curve distally, and the teeth decrease rapidly in size. The granular 
ridge on the upper surface of the manus is about the same in both species, but is 
slightly less prominent in P. marginatus than P. dissimulatus. 

Beginning with Stimpson (1859), many authors (Young 1900; Chace 1956; Haig 
1956; Holthuis and Manning 1970; Gore 1974; Werding 1977) have alluded to the 
often striking hues of purple, crimson or Van Dyke red in P. marginatus (and as 
P. cessacii). Unfortunately, there are no references to color for P. dissimulatus 
other than Benedict’s or Schmitt’s brief notes mentioned above. At present it 
seems likely that P. dissimulatus is probably red, with banded red and white 
walking legs, and thus slightly different in color from P. marginatus. 

The two species are also distinguishable in size. Recall that Stimpson’s spec- 
imen was about 12 mm cl, and P. marginatus can grow to nearly 18 mm cl (based 
on a molted carapace from Isla Aves, Venezuela; Manning in litt.). Ovigerous 
females range from 7—12.6 mm cl in West Africa (Chace 1956; Holthuis and 
Manning 1970), and 9.5—17.5 mm cl in the Caribbean region (Gore 1974, unpub- 
lished data). On the other hand, P. dissimulatus apparently does not grow larger 
than about 6.5 mm cl, and ovigerous females range from 3.9-5.7 mm cl, so that 
the species is clearly smaller than P. marginatus. 


100 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Finally, P. dissimulatus is quickly distinguished not only from P. marginatus, 
but from all other porcellanids in the western Atlantic Ocean with the exception 
of Petrolisthes rosariensis Werding, 1977, by the presence of double epibranchial 
spines. In P. marginatus an extremely small nub of a second spine, more like an 
enlarged tubercle than a spine or spinule, may occasionally occur behind one 
epibranchial spine or the other, but this remains the exception not the rule. More- 
over, P. rosariensis is easily separated because its chelipeds and carapace are 
covered with transverse piliferous rugae much as in P. galathinus, to which it is 
closely related. Neither P. dissimulatus nor P. marginatus has transverse pili- 
ferous rugae anywhere on the carapace or chelipeds. 


Acknowledgments 


I thank Drs. Raymond B. Manning and Fenner A. Chace, Jr., National Museum 
of Natural History, Washington, D.C. for alerting me to this problem and pro- 
viding literature sources, and much of their own data. I also thank the following 
colleagues for searching their respective collections for specimens of the three 
species considered in this report: Dr. Jacques Forest, Muséum National d’His- 
toire Naturelle, Paris (MNHN); Janet Haig, Allan Hancock Foundation, Univer- 
sity of Southern California (AHF); Williard D. Hartmann, Yale Peabody 
Museum, New Haven (YPM); Bernd Hauser, Muséum d Histoire Naturelle de 
Geneve (MHNG); Lipke B. Holthuis, Rijksmuseum van Natuurlijke Historie, 
Leiden (RMNH); R. W. Ingle, British Museum (Natural History), London 
(BMNH); Herbert Levi, Museum of Comparative Zoology, Harvard (MCZ); Ray- 
mond B. Manning (USNM); A. Solem, Field Museum of Natural History, Chi- 
cago (FMNH); Gilbert L. Voss, University of Miami (UMML); and Torben Wolff, 
Universitets Zoologiske Museum, Copenhagen (UZMC). Dr. George Davis, 
Academy of Natural Sciences, Philadelphia (ANSP) graciously provided access 
to stored collections, and Dr. Bernd Werding, Institut flr Allgemeine und Spe- 
zielle Zoologie, Giessen, generously loaned specimens of the new species and P. 
marginatus from his private collection. 


Literature Cited 


Balss, H. 1914. Decapode Crustaceen von den Guinea-Inseln, Stid-Kamerun und dem Congoge- 
biet.—Ergibnisse der Zweiten Deutschen Zentral-Afrika-Expedition 1910-1911 unter Fuhrung 
Adolph Friedrichs, Herzog zu Mecklenburg 1, Zoologie:97—108, figs. 1-12. 

—. 1922. Crustacea VII: Decapoda Brachyura (Oxyrhyncha und Brachyrhyncha) und geogra- 
phische Ubersicht tiber Crustacea Decapoda.In W. Michaelsen, Beitrage zur Kenntnis der 
Meeresfauna Westafrikas 3(3):69-1 10. 

Benedict, J. E. 1901. The Anomuran collections made by the Fish Hawk expedition to Porto Rico.— 
Bulletin of the U.S. Fish Commission 20(2):129-148, 3 figs., pls. 4—6. 

Boschma, H. 1931. Papers from Dr. Th. Mortensen’s Pacific Expedition 1914-16. LV. Rhizoceph- 
ala.—Videnskabelige Meddelelser fra Dansk Naturhistorisk Forening i Kjgbenhavn 89:297— 
380, text-figs. 1-57. 

Chace, F. A., Jr. 1956. Porcellanid crabs.—Expédition Océanographique Belge dans les eaux co- 
tieres Africaines de |’ Atlantique Sud (1948-1949), Résultats Scientifiques 3(5):1—54, figs. 1-14. 

——. 1962. The non-brachyuran decapod crustaceans of Clipperton Island.—Proceedings of the 
United States National Museum 113(3466):605—635, figs. 1-7. 

Coelho, P. A. 1966. Lista dos Porcellanidae (Crustacea, Décapoda. Anomura) do Litoral de Per- 
nambuco e dos Estado Vizinhos.—Trabajos de la Instituto Oceanografico de Universidade do 
Recife 5/6:51-68. 


VOLUME 96, NUMBER 1 101 


——. 1970. A distribuigao dos Crustaceos Decapodos Reptantes do norte do Brasil.—Trabajos 
de la Instituto Oceanografico de Universidade Federal de Pernambuco, Recife 9/1 1:223-238, 
fig. 1. 

, and M. de Araujo Ramos. 1972. A constituigao e a distribuigao de Fauna de Decapodos 

do litoral leste da América do Sul entre as Latitudes de 5° N e 59° S.—Trabajos de la Instituto 

Oceanografico de Universidade Federal de Pernambuco, Recife 13:133—236, figs. 1-4. 

Evans, A. C. 1967. Syntypes of Decapoda described by William Stimpson and James Dana in the 
collections of the British Museum (Natural History).—Journal of Natural History 1(3):399- 
411. 

Fausto Filho, J. 1974. Stomatopod and decapod crustaceans of the Archipelago of Fernando de 
Noronha, Northeast Brazil.—Arquivos de Ciéncias do Mar (Fortaleza) 14(1):1-35. 

Gauld, D. T. 1960. Decapoda Anomura. An annotated check list of the Crustacea of Ghana. IIJ.— 
Journal of the West African Scientific Association 6(1):64—67. 

Gore, R. H. 1974. Biological results of the University of Miami Deep-Sea expeditions, 102. On a 
small collection of porcellanid crabs from the Caribbean Sea (Crustacea, Decapoda. Anomu- 
ra).—Bulletin of Marine Science 24(3):700-721, figs. 1-5. 

—. 1982. Porcellanid crabs from the Pacific and Atlantic coasts of Central America (Crustacea 
Decapoda: Anomura).—Smithsonian Contributions to Zoology 363:i-1v, 1-34. 

Haig, J. 1956. The Galatheidea (Crustacea Anomura) of the Allan Hancock Atlantic Expedition 
with a review of the Porcellanidae of the western North Atlantic.—Allan Hancock Atlantic 
Expedition 8:1—43, | pl. 

—. 1962. Papers from Dr. Th. Mortensen’s Pacific Expedition 1914-1916, LX XIX: Porcellanid 

crabs from eastern and western America.—Videnskabelige Meddelelser fra Dansk Naturhis- 

torisk Forening 1 Kjgbenhavn 124:171—192, figs. 1-5. 

1964. Papers from Dr. Th. Mortensen’s Pacific Expedition 1914-1916. 81. Porcellanid crabs 
from the Indo-West Pacific, Part |.—Videnskabelige Meddelser fra Dansk Naturhistorisk For- 
ening i Kjgbenhavn 126:355-386, figs. 1-4. 

—. 1974. The anomuran crabs of Western Australia: Their distribution in the Indian Ocean and 
adjacent seas.—Journal of the Marine Biological Association of India 14(2):443-451. 

Holthuis, L. B., and R. B. Manning. 1980. The R/V Pillsbury Deep-Sea Biological Expedition to 
the Gulf of Guinea, 1964-65. 12. The Porcellanidae, Hippidae, and Albuneidae (Crustacea, 
Decapoda).—Studies in Tropical Oceanography 4(2):241—255, fig. 1. 

Lebour, M. V. 1959. The larval decapod Crustacea of tropical West Africa.—Atlantide Report 5: 
119-143, figs. 1-21. 

Miers, E. J. 1881. Crustacea. Jn Report on a collection made by Mr. T. Conry in Ascension Island.— 
Annals and Magazine of Natural History, (5) 8:432—434. 

Milne Edwards, A. 1878. Description de quelques espéces nouvelles de Crustacés provenant du 

voyage aux iles du Cap-Vert de M. M. Bouvier et de Cessac.—Bulletin de la Société Philom- 

athique de Paris, (7):225—232. 

, and E. L. Bouvier. 1900. Crustacés Décapodes. I. Brachyures et Anomoures. Premiere 

partie.—Expédition scientifiques du Travailleur et due Talisman pendant les Années 1880, 

1881 1882, et 1883. 396 pp., 32 pls. Paris: Masson. 

Nobili, G. 1897. Decapodi et Stomatopodi raccolti dal Dr. Enrico Festa nel Darien a Curagao, La 
Guayra, Porto Cabello, Colon, Panama, ecc.—Bollettino Musei di Zoologia ed Anatomia Com- 
parata della R. Universita di Torino 12(280):1-8. 

—. 1901. Viaggio del Dr. Enrico Festa nella Repubblica dell’Ecuador e regioni vicine, 23: 
Decapodi e Stomatopodi.—Bollettino Musei di Zoologia ed Anatomia Comparata dell R. Univ- 
ersita di Torino 16(415):1—58. 

Pocock, R. I. 1890. Crustacea. In Ridley, Notes on the zoology of Fernando Noronha.—Journal 
of the Linnean Society of London, Zoology 20:506—526. 

Rickner, J. A. 1975. Notes on members of the family Porcellanidae (Crustacea: Anomura) collected 
on the east coast of Mexico.—Proceedings of the Biological Society of Washington 88(16):159— 
166. 

Schmitt, W. L. 1924a. The macruran, anomuran and stomatopod Crustacea. Bijdragen tot de kennis 
der fauna van Curacao.—Resultaten eener reis van Dr. C. J. Van Der Horst in 1920, 23:61— 
81, figs. 1-7, pl. VIII. 

——. 1924b. Report on the Macrura, Anomura and Stomatopoda collected by the Barbados- 


102 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Antigua Expedition from the University of Iowa in 1918.—University of Iowa Studies in Nat- 
ural History 10(4):65—99, pls. I-IV. 

——., 1935. Crustacea Macrura and Anomura of Porto Rico and the ‘Virgin Islands.—Scientific 
Survey of Porto Rico and the Virgin Islands, New York Academy of Sciences 15(2):125—227, 
figs. 1-80, pls. LIV. 

Sourie, R. 1954. Contribution a l’étude écologique des cétes rocheuses du Sénégal.—Mémoirs de 
l'Institut Frangais d’ Afrique Noire 38:1-342, pls. 1-23. 

Stimpson, W. 1858. Prodromus descriptionis animalium evertebratorum, quae in expeditione ad 
Oceanum Pacificum Septentrionalem, a Repubblica Federata Missa, Cadwaladaro Ringgold 
et Johanne Rodgers ducibus, observavit et descriptsit, pars VII, Crustacea Anomura, I: Te- 
leosomi.—Proceedings of the Academy of Natural Sciences of Philadelphia (1859) 10:225—252. 

——. 1859. Notes on North American Crustacea, No. I.—Annals of the Lyceum of Natural 
History in New York 7:49-93, 1 pl. 

Werding, B. 1977. Los Porcelanidos (Crustace: Anomura: Porcellanidae) de la region de Santa 
Marta, Colombia.—Annales de Instituto de Investigaciones Marinas de Punta de Betin 9:173-— 
214, figs. 1-29. 

Young, C. G. 1900. The stalk-eyed Crustacea of British Guiana, West Indies, and Bermuda. John 
M. Watkins, London. 514 pp., 7 pls. 


66-1 Drexelbrook Dr., Drexel Hill, Pennsylvania 19026. 


PROC. BIOL. SOC. WASH. 
96(1), 1983, pp. 103-109 


GEOGRAPHIC VARIATION IN 
CHLOROSPINGUS OPHTHALMICUS IN COLOMBIA 
AND VENEZUELA (AVES: THRAUPIDAE) 


Storrs L. Olson 


Abstract.—Geographic variation in the montane species Chlorospingus 
ophthalmicus was studied in specimens from the Andes of Colombia and Vene- 
zuela. There are two distinct subspecies groups in this area that were formerly 
treated as separate species. The white postocular spot is lacking in the flavopectus 
group, which has five representatives in Colombia, including two described as 
new (C. o. trudis n. subsp. and C. o. exitelus n. subsp.). Additional taxa are 
found in Ecuador and Peru. In the ophthalmicus group there are five named 
subspecies in Colombia and Venezuela, as well as two populations of intergrades 
or undetermined status. In this group, the subspecies C. 0. jacqueti has three 
curiously disjunct populations separated by distinct intervening forms. Specimens 
of the flavopectus group are reported from a locality only 40 km from the nearest 
known population of the ophthalmicus group, which suggests the possibility of 
determining whether two distinct species are actually involved. Further field work 
and collecting are needed to resolve some of the intriguing problems in the evo- 
lutionary history of C. ophthalmicus in Colombia and Venezuela. 


The Common Bush-Tanager, Chlorospingus ophthalmicus, exhibits complex 
variation through its extensive range from Mexico through Argentina. A large 
series of specimens from Colombia in the National Museum of Natural History, 
Smithsonian Institution (USNM), most of which had not previously been iden- 
tified to subspecies, prompted the following revision. To effect this, I assembled 
material from other institutions, including representation from Venezuelan pop- 
ulations. Although I have included my observations on the Venezuelan speci- 
mens, these are tentative and more material will have to be consulted before the 
systematics and distribution of some of the forms of C. ophthalmicus are properly 
understood in that country. The spelling of Colombian place names, and their 
location by department, conforms with that of Paynter and Traylor (1981) and is 
not necessarily that found on specimen labels or in my previous publications. 
The distribution of the various populations of C. ophthalmicus in Colombia and 
Venezuela is shown in Fig. 1. 


Chlorospingus ophthalmicus jacqueti Hellmayr, 1921 


Type-locality.—Galipan, Cerro del Avila, near Caracas, Venezuela. 

Characters.—White postocular spot and dark throat speckles present. Throat 
only slightly tinged with buffy. Crown color relatively light, rather grayish brown. 
Yellow breast-band only moderately suffused with orangish. 

Range.—Inexplicably disjunct. Found in northern Venezuela from Miranda 
west to Carabobo, then in Trujillo (and extreme eastern Mérida according to 


104 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


CHLOROSPINGUS OPHTHALMICUS 
@ Jacquet! 


BX) sussp. 
© FALCONENSIS 


@ Venezuet anus 


KK Pons! 

© EMINENS 

© vacaueti > EMINENS 
A FLAvoPectus 

@ TRUDIS 

@ MACARENAE 

MM Nicriceps 

L\ EXITELUS 


Fig. 1. Distribution of Chlorospingus ophthalmicus in Colombia and Venezuela based on material 
examined in this study. Some specimens (AMNH and MCZ) were examined after the map had been 
prepared and a few localities are therefore not shown here. Department boundaries in Colombia do 
not reflect several of the more recent shifts and splits. Hatching indicates areas above 1000 feet 
elevation. 


Storer [1970]), and finally on the western slope of the eastern Andes of Colombia 
in southern Cesar, Norte de Santander, and northern Santander. 

Specimens examined.—VENEZUELA. MIRANDA: Pico de Naiguata (1, 
USNM; 1, CM). DISTRITO FEDERAL: El Limon (4, CM). ARAGUA: Cordi- 
llera de la Costa, Rancho Grande (7, USNM; 1, LSU); Colonia Tovar (5, CM; 
1, USNM). CARABOBO: La Cumbre de Valencia (8, CM). TRUJILLO: Guamito 
(10, CM; 2, MCZ). 

COLOMBIA. CESAR: La Palmita (22, CM). NORTE DE SANTANDER: Ra- 
mirez (2, CM); Las Ventanas (7, CM). SANTANDER: Corcova [not in Paynter 
and Traylor] (2, LACM); Cachiri (2, FM); Hacienda Las Vegas (1, USNM). 

Remarks.—Although the distribution of this subspecies makes no sense, I am 
forced to concur with Zimmer (1946, 1947) that the birds from the west slope of 
the northern part of the eastern Andes of Colombia are inseparable from Vene- 
zuelan jacqueti. These birds are separated from those of Trujillo by the very 
different subspecies eminens and venezuelanus, and those of Trujillo are in turn 
separated from those of northern Venezuela by an unnamed dark-capped form 


VOLUME 96, NUMBER 1 105 


from Lara and the very distinctive race falconensis of Falc6n and Yaracuy. K. 
C. Parkes (in litt. 12 July 1982) notes that there are slight differences in coloration 
of the dorsum and undertail coverts between the three disjunct populations of 
jacqueti. Nevertheless, these populations are clearly much more similar to one 
another than to any of the intervening forms. 


Chlorospingus ophthalmicus subsp. indet. 


Characters.—Near jacqueti but crown noticeably darker, less grayish, though 
not as dark as in venezuelanus; buffiness of throat and orangish of breast band 
not as marked as in venezuelanus or falconensis. 

Range.—Venezuela, in Lara and northern Trujillo. 

Specimens examined.—VENEZUELA. LARA: Anzoategui (3, CM); Guarico 
(4, CM). TRUJILLO: Paramo de Rosas (2, CM). 

Remarks.—The darker crown in specimens from Lara and Paramo de Rosas 
has already been noted by Hellmayr (1936:403, footnote). It is consistent and 
prevents these birds from being assigned to jacqueti. They could perhaps be 
considered as intergrades with venezuelanus, as Hellmayr was inclined to do, but 
the intervening population of apparently typical jacqueti from Guamito, Trujillo, 
complicates such an interpretation. Specimens of C. ophthalmicus from a number 
of additional localities from Mérida through Yuracuy would be desirable. 


Chlorospingus ophthalmicus falconensis Phelps and Gilliard, 1941 


Type-locality.—San Luis Mountains, above San Luis, Falcon, Venezuela. 

Characters.—Crown much darker than in jacqueti, but browner and not as 
dark blackish as in venezuelanus. Throat buffier and breast band more orangish 
than in either jacqueti or venezuelanus. 

Range.—Venezuela, in the states of Falcon and Yaracuy. 

Specimens examined.—VENEZUELA. FALCON: San Luis (1, USNM). 
YARACUY: Lagunita de Aroa (1, CM). 


Chlorospingus ophthalmicus venezuelanus Berlepsch, 1893 


Type-locality.—Mérida, Venezuela. 

Characters.—Crown much darker than in any of the other subspecies having 
a white postocular spot, with the exception of eminens. Differs from eminens 
(and jacqueti) in the buffier throat and more orangish breast band. 

Range.—Southwestern Venezuela in the Andes of Mérida, Tachira, and por- 
tions of Lara (according to Storer [1970], no specimens from Lara or Tachira 
seen). 

Specimens examined.—VENEZUELA. MERIDA: Culata (2, USNM); Escor- 
ial (1, ANSP); Heights Tabay (4, CM). 


Chlorospingus ophthalmicus ponsi Phelps and Phelps, 1952 


Type-locality.—Cerro Tamuypejocha, Sierra Perija, Zulia, Venezuela. 

Characters.—Like jacqueti but throat decidedly buffier, with more pronounced 
speckling, breast band more orangish. Crown much lighter than in venezuelanus 
or falconensis. Not as richly colored ventrally as falconensis. 


106 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Range.—The Sierra de Perija in Zulia, Venezuela, and Cesar and La Guajira, 
Colombia. 

Specimens examined.—COLOMBIA. CESAR: Hiroca (10, USNM). LA GUA- 
JIRA: La Africa (3, USNM); Tierra Nueva (8, USNM); Monte Elias (2, USNM). 

Remarks.—Hitherto this subspecies was known only from the vicinity of the 
type-locality, the above specimens being the first recorded for Colombia. They 
match the original description of ponsi perfectly except that I do not find the 
crown to be any darker than in jacqueti. There is considerable individual variation 
in the darkness of the crown in both subspecies, however. 


Chlorospingus ophthalmicus eminens Zimmer, 1946 


Type-locality.—Gramalote, Norte de Santander, Colombia. 

Characters.—Very similar to venezuelanus in the dark crown, differing only 
in the less buffy, nearly white throat and less orangish breast band. 

Range.—Colombia, on the eastern slope of the eastern Andes in southern Norte 
de Santander and Boyaca. 

Specimens examined.—COLOMBIA. NORTE DE SANTANDER: Gramalote 
(1, ANSP); Palo Gordo (3, USNM). BOYACA: Hacienda La Primavera (4, FM). 

Remarks.—Apparently intergrades with the westernmost of the three disjunct 
populations of jacqueti (see following account). 


Chloropsingus ophthalmicus jacqueti > eminens 


Characters.—Nearest jacqueti but crown darker, more blackish brown, less 
gray. Crown not as dark as in eminens, throat buffier and breast band more 
orangish. 

Range.—Colombia in Norte de Santander, geographically intermediate be- 
tween eminens to the east and jacqueti to the west. 

Specimens examined.—COLOMBIA: NORTE DE SANTANDER: Buenos 
Aires (8, USNM); Alto del Pozo (1, USNM); 8 mi. S of Convencion, (3, USNM); 
Ocana, beyond Pueblo Nuevo (1, USNM). 

Remarks.—The above series is quite distinct from either jacqueti or eminens 
and shows little variation, so that it would be possible to designate it as a new 
subspecies. I believe that at present, however, these birds should be regarded as 
intergrades. They are very similar to the unnamed dark-capped birds from Lara, 
Venezuela, and differ only in their somewhat buffier throats. 


Chlorospingus ophthalmicus flavopectus (Lafresnaye, 1840) 


Type-locality.—Santa Fé de Bogota, Colombia. 

Characters.—Differs from all of the preceding subspecies in lacking the white 
postocular spot; crown gray without any brownish cast; throat white, nearly 
without speckles; size larger. 

Range.—Colombia, on the western slope of the eastern Andes from Santander 
through Cundinamarca. 

Specimens examined.—COLOMBIA. SANTANDER: Above Virolin (4, 
USNM). BOYACA: Paramo de Agtiero (1, USNM); Arcabuca, 3 km SW (1, 
MVZ); Lago de Tuquina [not in Paynter and Traylor] (1, FM). CUNDINAMAR- 
CA: La Aguadita (2, ANSP); Subia (2, AMNH); Aguabonita (2, AMNH); El 


VOLUME 96, NUMBER 1 107 


Roble (2, AMNH); “Santa Fe de Bogota’ (1, USNM); ‘‘Bogota’’ (1, MCZ [ho- 
lotype]; 4, USNM). 

Remarks.—Hellmayr (1936) and earlier authors considered flavopectus, along 
with certain forms from Ecuador and Peru, to be specifically distinct from 
ophthalmicus. It was not until Zimmer’s (1947) revision that the two species were 
combined. The forms of the flavopectus group (which in Colombia includes fla- 
vopectus, trudis, macarenae, nigriceps, and exitelus) seem so distinct from those 
in the ophthalmicus group that I have reservations about Zimmer’s action and I 
wonder if the earlier treatment may not have been correct after all. There is a 
possibility that further collecting and field work in Santander could resolve this 
(see following account). 


Chlorospingus ophthalmicus trudis, new subspecies 


Holotype.—CM 59494, male, La Pica, Santander, Colombia. Collected 12 Feb- 
ruary 1917 by M. A. Carriker, Jr., original number 21063. 

Characters.—Most similar to flavopectus but dorsum lighter, more yellowish- 
green, less olivaceous; crown lighter, more brownish gray; auriculars brownish 
rather than black; inner webs of rectrices not blackish but brownish-gray or 
greenish. Averages smaller, with a visibly smaller bill (wing chord 64.8—72.1 mm, 
average 68.6, n = 8; vs. 67.4-76.4, average 72.3, n = 9, in flavopectus [72.2 mm 
in holotype]). 

Range.—Known so far only from the type-locality. Meyer de Schauensee (1948: 
320) describes La Pica as a campsite at 2800 m on “‘the stretch of mountain trail 
crossing the ridge known as Cruz de Piedra, between San Andrés and Malaga, 
north of Molagavita.”’ 

Specimens examined.—Holotype and 8 topotypes with essentially the same 
data (6, CM; 2, ANSP; 1, MCZ). 

Etymology.—Latin trudis, a pointed pole or pike, one of several meanings for 
the Spanish word pica, in reference to the type-locality. 

Remarks.—This form is removed about 85 km to the north of the nearest 
specimens of flavopectus that I examined (near Virolin). Differentiation may have 
been affected by the barrier afforded by the valley of the Rio Chicamoca. Of 
more significance is the fact that less than 40 km separate trudis from populations 
of jacqueti to the north (at Hacienda Las Vegas) and there is no such barrier 
evident between them. It would be of great interest to determine how these two 
completely different forms interact if and where they come in contact. 


Chlorospingus ophthalmicus macarenae Zimmer, 1947 


Type-locality.—Mt. Macarena, Meta, Colombia. 

Characters.—Like flavopectus but smaller, throat slightly buffier, yellow breast 
band more restricted and more intensely colored, greenish flanks more extensive 
and yellower, belly white not grayish. 

Range.—Colombia, confined to the Macarena Mountains east of the eastern 
Andes in Meta. 

Specimens examined.—COLOMBIA. META: La Macarena (12, FM). 

Remarks.—This is a well marked isolate obviously allied with the flavopectus 
group. 


108 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Chlorospingus ophthalmicus nigriceps Chapman, 1912 


Type-locality.—Miraflores, 6800 ft., Central Andes, east of Palmira, Cauca, 
Colombia. 

Characters.—Similar to flavopectus in lacking the white postocular spot but 
crown very dark, blackish; throat and sometimes part of pectoral band heavily 
speckled with blackish; green of dorsum darker, suffused with blackish. 

Range.—Colombia, western slope of the southern end of the eastern Andes 
and both slopes of the southern end of the central Andes. 

Specimens examined.—COLOMBIA. QUINDIO: Laguneta (7, ANSP). HUI- 
LA: Moscopan (2, CM; |, FM); Tiyeras, Moscopan (5, USNM; 3, FM; 2, MVZ); 
La Plata (1, ANSP); Buenavista (6, ANSP); La Candela (16, ANSP; 1, AMNH); 
Andalucia (1, USNM). TOLIMA: Toche (5, ANSP; 1, MCZ); Rio Toche (1, 
USNM; 1, MCZ); El Edén (1, AMNH). VALLE DEL CAUCA: E. of Palmira 
(1, AMNH, holotype). 

Remarks.—Previously published records from the northern part of the central 
Andes in Antioquia pertain to the following new subspecies. 


Chlorospingus ophthalmicus exitelus, new subspecies 


Holotype.—USNM 403747, female, ovary enlarged. Ventanas, Valdivia, An- 
tioquia, Colombia, 7000-7400 feet. Collected 11 June 1948 by M. A. Carriker, 
Jr., original number 13878. 

Characters.—Nearest nigriceps but crown decidedly lighter, grayish instead of 
black; speckling on throat reduced and lighter in color; green of dorsum lighter, 
more yellowish. Crown darker than in flavopectus, in which there is almost no 
speckling on the throat. 

Range.—Colombia, both slopes of the northern end of the central Andes in 
Antioquia. 

Specimens examined.—COLOMBIA. ANTIOQUIA: Ventanas, Valdivia (3, 
USNM, including holotype); Hacienda Zulaiba (2, USNM); Envigado (1, USNM); 
Sabanalarga [not the Sabanalarga in Atlantico in Paynter and Traylor] (1, AMNH, 
Niceforo, collector); ““Medellin’? (1, AMNH); “‘Antioquia’’ [Salmon, collector] 
(1, MCZ); Santa Elena (6, AMNH, intergrades with nigriceps). 

Etymology.—Greek exitelos, a lessening or fading, in reference to the dimi- 
nution in intensity of the features that are characteristic of nigriceps. 

Remarks.—This subspecies is easily separated from nigriceps to the south. The 
birds from Sabanalarga and Medellin mentioned by Meyer de Schauensee (1951: 
1062) under nigriceps are definitely referable to this subspecies, as probably are 
those from Retiro (not seen). There apparently is but one record of C. ophthal- 
micus in the western Andes. Meyer de Schauensee (1951:1062) lists C. 0. nigri- 
ceps from Concordia, an old Salmon collecting site. This is presumably based on 
the record of “‘C. flavipectus’’ from Concordia mentioned in Sclater and Salvin 
(1879). As this is near the narrowest part of the Cauca Valley, with the foothills 
of the Central Andes being in places only a few hundred meters from the steep 
eastern slopes of the western Andes (W. L. Brown, in litt. 21 July 1982), dispersal 
of C. ophthalmicus from one range to the other at this point is not unlikely. The 
birds from the western Andes would almost certainly pertain to C. o. exitelus. 

A series of six specimens from Santa Elena, Antioquia, is quite variable, some 


VOLUME 96, NUMBER 1 109 


of the individuals having dark crowns, while others have lighter crowns. This 
series appear to be composed of intergrades between exitelus and nigriceps. None 
has the crown as dark as in nigriceps, however, but the throat is more heavily 
speckled than in exitelus. 


Acknowledgments 


For lending specimens crucial to this study I am indebted to Frank B. Gill, 
Academy of Natural Sciences of Philadelphia (ANSP); Kenneth C. Parkes, Car- 
negie Museum of Natural History, Pittsburgh (CM); John W. Fitzpatrick and 
Melvin A. Traylor, Jr., Field Museum of Natural History, Chicago (FM); John 
P. O'Neill, Louisiana State University, Baton Rouge (LSU); and Ned K. John- 
son, Museum of Vertebrate Zoology, University of California, Berkeley (MVZ). 
I also examined specimens in the collections of the American Museum of Natural 
History, New York (AMNH), and the Museum of Comparative Zoology, Harvard 
University (MCZ). For reading and commenting on the manuscript I am grateful 
to W. L. Brown, S. L. Hilty, Kenneth C. Parkes, and Raymond A. Paynter, Jr. 
This is contribution number |2 of the Wetmore Papers, a project supported in 
part by trust funds from the Smithsonian Institution for completing unfinished 
work and study of undescribed material left by the late Alexander Wetmore. 
Janine Higgins prepared the map. 


Literature Cited 


Hellmayr, C. E. 1936. Catalogue of Birds of the Americas. Part 9.—Field Museum of Natural 
History, Zoological Series 13(9):1—458. 

Meyer de Schauensee, R. 1948. The birds of the Republic of Colombia. Part 1.—Caldasia 5:251- 

380. 

1951. The birds of the Republic of Colombia. Part 4.—Caldasia 5:873-1112. 

Paynter, R. A., Jr., and M. A. Traylor, Jr. 1981. Ornithological gazetteer of Colombia.—Harvard 
University, Cambridge, Massachusetts. 

Sclater, P. L., and O. Salvin. 1879. On the birds collected by the late Mr. T. K. Salmon in the State 
of Antioquia, United States of Columbia.—Proceedings of the Zoological Society of London 
1879:486—550. 

Storer, R. W. 1970. Subfamily Thraupinae. Pp. 246-408 in R. A. Paynter, Jr., editor. Check-list of 
Birds of the World. Vol. 13.—Museum of Comparative Zoology, Cambridge, Massachusetts. 

Zimmer, J. T. 1946. A new subspecies of tanager from northeastern Colombia.—Journal of the 
Washington Academy of Sciences 36(11):389-390. 

———. 1947. Studies of Peruvian birds. No. 52.—American Museum Novitates 1367:1—26. 


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


PROC. BIOL. SOC. WASH. 
96(1), 1983, pp. 110-114 


FUNCTION OF THE TEETH AND VESTIBULAR ORGAN 
IN THE CHAETOGNATHA AS INDICATED BY 
SCANNING ELECTRON MICROSCOPE 
AND OTHER OBSERVATIONS 


Robert Bieri, Dolores Bonilla, and Fernando Arcos 


Abstract.—Ultrastructure of the teeth of chaetognaths indicates that they may 
function to pierce the exoskeleton of copepods. The vestibular ridge behind the 
posterior teeth has a series of pores that may excrete a poison that immobilizes 
copepods. 


A series of recently published SEM photographs of the hooks and teeth (ar- 
mature) of several species of chaetognaths (Cosper and Reeve 1970; Nagasawa 
and Marumo 1973, 1979; Bone and Pulsford 1978; Spero, Hagan, and Vastano 
1979) permits comparison with some SEM photographs we have made. The an- 
terior and posterior teeth have a distinct microstructure at the tips reminiscent 
of a stone drill. We postulate that they puncture copepod exoskeletons to speed 
the penetration of digestive enzymes and hasten absorption of digested material. 
The teeth also have lightly serrate, knife-like edges and thus also can cut or shear. 

A series of pores in the papillae of the vestibular organ, visible in earlier SEM 
studies may be the openings of ducts for the secretion of a toxin. The toxin could 
flow down the grooves of the teeth and penetrate the prey through the puncture 
holes. If these hypotheses are confirmed, then the epithet, “‘Arrowworms, Tigers 
of the Sea’’ should be changed to *‘Arrowworms, Cobras of the Sea.”’ 

Methods.—Each study published to date has used different methods of killing, 
preservation, and sample manipulation. The methods of Bone and Pulsford (1978) 
and of Spero, Hagan, and Vastano (1979) have given the finest pictures of soft 
tissue. We used both formalin and dilute (0.4%) gluteraldehyde for killing and 
fixing. The latter gave better preservation of the soft tissues, but where live 
animals are not available, formalin killed specimens can give useful information. 
We used both acetone and isoamylacetate for final dehydration and found that 
the armature of chaetognaths is sufficiently rugged that we could dry specimens 
from acetone using a heat lamp and still get good gold plating. 

Results.—Spero, Hagan, and Vastano very generously allow us to publish here 
their previously unpublished, superb SEM picture of the head of Sagitta hispida 
(Fig. 1). This is useful for orientation and shows the microstructure of the teeth 
(Fig. 1). The hooks or seizing jaws are on either side of the head. The two paired 
rows of smaller anterior teeth and larger posterior teeth are clearly visible, as are 
the papillae along the ridge of the vestibular organs under each set of posterior 
teeth. For comparison, Fig. 2 shows the head of Prerosagitta draco. The muscular 
‘‘hood”’ is partly drawn over the hooks on the right side of the head. Five, 
possibly six anterior teeth on the right side of the head and six or seven on the 
other side are visible but the posterior teeth are too numerous to count accurately. 
The velvet-like tissue around the mouth, clearly shown in Fig. 1, is badly con- 


VOLUME 96, NUMBER 1 111 


Fig. 1. SEM of the head of Sagitta seen from the front and below. The hooks (also called seizing 
jaws and spines) are arranged on either side of the head. The anterior and posterior teeth are arranged 
in two pairs of rows, the anterior teeth projecting diagonally and the posterior teeth hanging down in 
this preparation. Photo courtesy of Spero. Hagan, and Vastano. 


tracted either by the formalin fixation or by the dehydration. The vestibular organ 
is strongly wrinkled and contracted and shows up as a series of white knobs on 
narrow stalks. The knobs are shown enlarged in Fig. 3. This severe distortion of 
the soft tissues proved fortuitous because it strongly emphasizes the pores present 
in each papilla of the vestibular organ. Refering back to Fig. 1, it is barely possible 
to see that they are present also in Sagitta hispida. We also found them present 
in Sagitta peruviana (Fig. 4). Here the pores are clearly evident although it is 
not clear if they are present in every papilla as they appear to be in Prerosagitta 
draco. The pores, though small, are shown clearly in Fig. | of Sagitta hispida in 
Cosper and Reeve (1970). 

The microstructure of the teeth seems to be essentially the same in all three 
species. Though somewhat blurred, the tip structure shows up best in Fig. 4. 
There are four cusps at the tip, one is longer and the other three are shorter and 
subequal in length. The tips of several different teeth are outlined in Fig. 5. All 
can be interpreted as different views of the 4-cuspate form. Close examination 
of Fig. 2 of Cosper and Reeve (1970) shows the same quadri-cuspate structure. 

Discussion.—The shape of the tips of the teeth seems well suited for quick 
penetration of the longest cusp into a rigid surface followed by a splitting or 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


VOLUME 96, NUMBER 1 113 


Fig. 5. Outlines of anterior and posterior teeth taken from SEM photos showing how the quad- 
ridentate pattern can appear as a single, bifid or trifid point. The larger sketches are of Sagitta 
peruviana, the smaller of Pterosagitta draco and are not drawn to scale. 


perforating of the exoskeleton by the three following cusps. The serrate, knife- 
like edges of the teeth could serve as slicing or cutting edges on soft-bodied 
animals such as fish larvae. Robert Miller (personal communication) stated that 
during his experimental studies on fish, unidentified sagittas cut many of the 
larvae cleanly in half. 

For the pores in the vestibular organ, three possible functions suggested them- 
selves. They could hold prey by suction. They could secrete digestive enzymes, 
or an immobilizing toxin. In reviewing the literature related to head structure, 
digestive physiology, and feeding behavior (summarized by Hyman 1959) we 
could find no indication of the internal structure of the vestibular organs. Parry 


— 


Fig. 2. Head of Pterosagitta draco showing hooks (partly covered by the hood on the right side 
of the head) anterior and posterior teeth, and the vestibular region highly distorted in preparation. 
Below the posterior teeth on the right side of the head the papillae of the vestibular organ appear as 
white knobs. 

Fig. 3. Enlargement of the same specimen as in Fig. 2 showing the vestibular ridge of the vestibular 
organ below the posterior teeth. Because of distortion the papillae appear as tubes. 


Fig. 4. Vestibular organ of Sagitta peruviana behind the posterior teeth. Pores in the vestibular 
ridge are clearly evident as is the cuspate or quadridentate shape of the teeth. 


114 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


(1947) found no toxin in the secretion from the anterior part of the pharynx. On 
the other hand, in describing the feeding behavior of Spadella which he found 
essentially the same as in Sagitta setosa, he wrote, “As the prey is clasped 
against the mouth region it is probably coated with secretion produced by the 
granular cells. By this means the appendages would be entangled and the copepod 
prevented from escaping. That the prey is not killed is shown by its circulating 
blood and gut peristalsis which can be observed for some time after swallowing. 
On the other hand, a copepod which has been relinquished is unable to move 
away. (Parry 1947:25). 

It seems to us that the vestibular organ could secrete a toxin specifically block- 
ing locomotion. Clearly this would be a great advantage to the arrowworm be- 
cause moving spines of even an herbivorous copepod could damage the mouth 
tissues. If the copepod’s circulation and gut movements continued to function as 
observed by Parry, it would speed the dispersion of the digestive enzymes. The 
teeth and hooks seem adequate to hold the prey against the mouth. The very 
close proximity of the pores of the vestibular ridge to the posterior teeth indicate 
a functional relationship between the two. The ridges of the posterior teeth, 
shown most clearly in Fig. 2 of Cosper and Reeve (1970) could direct a toxic 
secretion into the punctured body. 


Acknowledgments 


The authors extend sincere thanks to the Departamento de Microscopia Elec- 
tronica del Instituto Nacional de Hygiene, Guayaquil, for the use of their equip- 
ment and facilities. 


Literature Cited 


Bone, A., and A. Pulsford. 1978. The arrangement of ciliated sensory cells in Spadella (Chaetog- 
natha).—Journal of the Marine Biological Association of the United Kingdom 58:565-570. 

Cosper, T. C., and M. R. Reeve. 1970. Structural details of the mouthparts of a chaetognath as 
revealed by scanning electron microscopy.—Bulletin of Marine Science 20:441—445. 

Hyman, L. H. 1959. The invertebrates, 5, Smaller Coelomate groups. McGraw-Hill, New York, 
783 pp. 

Nagasawa, S., and R. Marumo. 1973. Structure of grasping spines of six chaetognath species 

observed by scanning electron microscopy.—Bulletin of the Plankton Society of Japan 19: 

63-75. 

, and 1979. Identification of chaetognaths based on morphological characteristics of 

hooks.—La Mer 17(4):14—24. 

Parry, D. A. 1947. Structure and function of the gut in Spadella cephaloptera and Sagitta setosa.— 
Journal of the Marine Biological Association of the United Kingdom 26:16—36. 

Spero. H., D. Hagan, and A. Vastano. 1979. An SEM examination of Sagitta tenuis (Chaeto- 
gnatha), utilizing a special sedation and handling procedure.—Transactions of the American 
Microscopical Society 98:139-141. 


(RB) Antioch College, Yellow Springs, Ohio 45389; (DB, FA) Instituto Ocean- 
ografico del Armado, Casilla 4950, Guayaquil, Ecuador. 


PROC. BIOL. SOC. WASH. 
96(1), 1983, pp. 115-120 


SYSTEMATICS AND DISTRIBUTION OF SHREWS OF 
THE GENUS CROCIDURA (MAMMALIA: 
INSECTIVORA) IN VIETNAM 


Lawrence R. Heaney and Robert M. Timm 


Abstract.—Three noncommensal species of white-tooth shrews (Crocidura) are 
found in Vietnam; from smallest to largest, they are C. horsfieldi indochinensis, 
C. attenuata, and C. fuliginosa dracula. All are probably widespread, although 
their distributions remain incompletely known in Vietnam. 


Shrews of the genus Crocidura are the most widespread and diverse insecti- 
vores in the Oriental faunal region (Chasen 1940; Ellerman and Morrison-Scott 
1951). However, there are few specimens from most areas, and this is especially 
true of the continental portions of southeast Asia. We recently surveyed the 
holdings of the American Museum of Natural History (AMNH), Field Museum 
of Natural History (FMNH), and U.S. National Museum of Natural History 
(USNM), and have encountered a number of misidentified specimens and im- 
portant unpublished records of three species of Crocidura from Vietnam. Because 
past investigators have been hampered by a lack of adequate descriptions and 
illustrations, we provide these here, along with reidentifications of previously 
published, misidentified specimens. Localities are mapped or listed in gazetteers 
by Moore and Tate (1965), Osgood (1932), and Van Peenen et al. (1969). We do 
not treat the common commensal species of the subgenus Suncus, C. murina, in 
this paper; see Van Peenen et al. (1969) for a list of localities for this species in 
southern Vietnam. 

All cranial measurements were taken by Heaney with dial calipers graduated 
to 0.05 mm. External measurements were taken from the collectors labels. The 
following cranial measurements were taken: condyloincisive length, condyles to 
anterior tip of incisors; braincase breadth, greatest breadth, roughly mastoidal; 
interorbital breadth, taken at anterior end of orbit; rostral length, anterior tip of 
incisor to anteriormost portion of orbit, in the infraorbital foramen; postpalatal 
depth, measured from just posterior to posterior lip of palate to closest point on 
cranial surface; rostral breadth, calipers held dorsally to rostrum, measurement 
taken at margin of P? and P?, in “‘notch’’; postpalatal length, taken from posterior 
edge of postpalatal lip to anteriormost point on foramen magnum; condyle to 
glenoid, calipers placed in glenoid fossa, drawn up against posteriormost point 
on occipital condyles; I‘ to M3, greatest length of toothrow, at alveolus of M? to 
anteriormost point on I‘; P* to M3, taken labially at alveolus; M? to M?, greatest 
width taken at anterior labial margins of second upper molars; palatal width at 
M3, alveolar distance between lingual margins of third upper molars. 


Crocidura attenuata Milne-Edwards 


Crocidura attenuata Milne-Edwards, 1872:263. 


These shrews are virtually identical to the species that follow in pelage char- 
acteristics; thus, the following description of the pelage applies equally well to 


116 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Table 1.—External and cranial measurements of white-toothed shrews from Vietnam. Values are 
means + standard deviation (and range). 


Measurements 


Crocidura attenuata 


(n = 


Crocidura fuliginosa 


(in = 17h) 


Crocidura horsfieldi 
(n = 4) 


Length of head and body 83.9 + 3.6 GBS se SY) 66.0 
(79-88) (83-101) (65-67) 
Length of tail 50.0 + 6.5 71.2 + 4.3 48.5 
(45-61) (62-80) (47-50) 
Length of hind foot 14.3 + 0.71 IO2 22 2 13.0 
(13-15) (15-18) (13) 
Condyloincisive length 20.88 + 0.95 23.63 + 0.43 17.77 
(19.6—22.4) (22.9-24.2) (17.7-17.9) 
Braincase breadth Yl 32 O27 10.23 + 0.18 8.13 
(9.1-9.9) (9.9-10.5) (8.0-8.2) 
Interorbital breadth 4.58 + 0.22 4.93 + 0.12 3.95 
(4.2-4.9) (4.7-5.1) (3.8-4.1) 
Rostral length 8.18 + 0.25 9.52 + 0.28 6.6 
(7.9-8.6) (8.8—9.8) (6.46.7) 
Postpalatal depth 3.77 + 0.14 4.18 + 0.13 3.48 
(3.5-3.9) (3.9-4.4) (3.4-3.6) 
Rostral breadth 2.67 + 0.19 2M a= Os iil 2.28 
(2.3-3.0) (2.6-2.9) (2.02.6) 
Postpalatal length 9.25 + 0.47 10.41 + 0.29 8.27 
(8.6—9.9) (10.0-11.0) (7.9-8.5) 
Condyle to glenoid 8.20 + 0.39 8.97 + 0.15 7.20 
(7.7-8.7) (8.89.2) (7.2) 
I’ to M3 9.08 + 0.36 10.65 + 0.21 7.48 
(8.69.7) (10.3-11.0) (7.3-7.6) 
P* to M4 5.09 + 0.21 5.99 + 0.15 4.20 
(4.95.5) (5.7—-6.2) (4.1-4.4) 
M2? to M? (labial) 6.39 + 0.38 6.95 + 0.21 5.35 
(5.7-6.8) (6.5—7.4) (4.9-5.7) 
Palatal width at M? 2.84 + 0.17 2.99 + 0.08 2.20 
(2.63.1) (2.8-3.1) (2.0-2.3) 


all three species. Dorsal pelage soft, dense; color grayish brown, hairs often 
tipped with silver, slate gray at base. Ventral pelage shorter, slightly more gray- 
ish. Ear covered inside and out with fine, scattered brown hairs; these hairs are 
densest along edge, giving ear a brownish fringe. Distal septum of ear with stiff 
brown hairs protruding beyond margin. Fore-feet and hind-feet with sparse cov- 
ering of brown hairs on dorsum; ventral surface naked of hairs. Tail thinly covered 
with brown hairs. Long hairs sparse but present on proximal two-thirds of tail, 
absent from distal third. Ventral surface of tail paler, grading into darker dorsum. 

Crocidura attenuata differs from C. fuliginosa externally in its generally small- 
er size (Table 1), and especially in its proportionately and absolutely shorter tail; 
the average tail to head and body ratio is 0.50. Crocidura attenuata differs ex- 
ternally from C. horsfieldi in being considerably larger (head and body more than 
79 mm as opposed to less than 67 mm), and in having a proportionately shorter 
tail. 


VOLUME 96, NUMBER 1 117 


Fig. 1. Dorsal, lateral, and ventral views of the cranium and lateral view of the mandible of 
Crocidura attenuata from Mt. Langbian, Tuyen Duc Province, Vietnam (FMNH 46640). Approxi- 
mately x4. 


The skull of C. attenuata (Figs. 1, 2) is similar to those of C. fuliginosa and 
C. horsfieldi in most respects. It differs from C. fuliginosa in being generally 
smaller; having a shorter and less massive toothrow, especially in having narrow- 
er molars (Fig. 2); having the second upper unicuspid conspicuously smaller than 
first or third; more rounded (U-shaped) base to the basisphenoid between the 
pterygoid processes; and a narrower strut of the maxillary over the infraorbital 
canal (Fig. 2). Crocidura attenuata differs from C. horsfieldi in having a larger 
skull (Table 1); a proportionately broader anterior tip to the rostrum; more pro- 
cumbent incisors; and in having the third unicuspid about three-fourths the size 
of the first, rather than about half (Fig. 2). 

These are the first specimens reported from Vietnam. As noted by Jenkins 
(1976), C. attenuata often has been confused with C. fuliginosa, and most of the 
specimens we examined have been reported previously in the literature as C. 
fuliginosa. Our identifications are based on measurements provided by Jenkins 
(1976), on comparisons with specimens (FMNH) from near the type locality of 
C. attenuata (see Allen 1938), and the illustrations and description published by 
Milne-Edwards (1872). Only one of the specimens from the USNM reported by 
Van Peenen et al. (1969) as C. fuliginosa is actually that species; the others are 
C. attenuata. The specimens cited here indicate that C. attenuata is widespread 
in southern Vietnam at elevations from near sea level to 1700 m. We note that 
the cranium of a Crocidura from Thailand figured by Lekagul and McNeely (1977: 
31) and identified as C. fuliginosa is actually C. attenuata, whereas the skull and 


118 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 2. Maxillary toothrows of Crocidura fuliginosa (A; FMNH 32422 from Chapa, Tonkin), C. 
attenuata (B; FMNH 46640 from Langbian Peak, Tuyen Duc Prov.), and C. horsfieldi (C; FMNH 
39029 from Chapa, Mt. Fan Si Pan, Tonkin); and infraorbital regions of C. fuliginosa (D; USNM 
357438 from Con Son Island), C. attenuata (E; FMNH 46641 from Langbian Peak, Tuyen Duc 
Prov.), and C. horsfieldi (F; USNM 357862 from An Khe Military Base, Binh Dinh Prov.). All 
drawings to same scale, approximately x9. 


measurements Lekagul and McNeely (1977:34) attribute to C. attenuata are those 
of C. fuliginosa. 

As noted by Jenkins (1976), the subspecific status of Indochinese specimens is 
uncertain, although Lekagul and McNeely (1977) thought Thai specimens prob- 
ably represented C. a. attenuata. 

Specimens examined.—VIETNAM: Quan Tri Prov.: 2.3 km E, 8 km S Thon 
Ke Tri Peak, elev. 70 m (1 USNM); Thua Thien Prov.: 8 km W, 3.7 km N Nui 
Ke, elev. 30 m (1 USNM); 9.1 km W, 3.6 km N Nui Ke, elev. 30 m (1 USNM); 
Tuyen Duc Prov.: Fyan, elev. 1200 m (1 USNM); Mt. Langbian (no elevation 
given; 3 FMNH); Mt. Langbian, elev. 1700 m (2 USNM). 


Crocidura fuliginosa dracula Thomas 


Crocidura fuliginosa dracula Thomas, 1912:686. 


As noted above, these shrews have pelage virtually identical to that of the 
other species, although we note that the skin on the hind feet of this species 


VOLUME 96, NUMBER 1 119 


appears to be paler dorsally than that of C. attenuata or C. horsfieldi. Crocidura 
fuliginosa differs cranially from C. attenuata as noted above; it differs from C. 
horsfieldi most conspicuously in its vastly larger size (Table |, Fig. 2), and also 
in its proportionately larger P*. 

Crocidura fuliginosa is widespread in southeast Asia (Jenkins 1976; Medway 
1977), and has been reported previously from Vietnam (Osgood 1932); most of 
the specimens we examined were reported by Osgood. As noted above, Van 
Peenen et al. (1969) reported this species from Con Son Island; their other records 
are here referred to C. attenuata. We have records of C. fuliginosa only from 
Con Son Island and from northern Vietnam, from elevations from 40 m to ca. 
1600 m, although the species is known to occur in southern Thailand and Malaysia 
(Lekagul and McNeely 1977; Medway 1977). This species apparently occurs sym- 
patrically with C. horsfieldi near Chapa in northwestern Tonkin. Van Peenen ef 
al. (1969) cited Thomas (1927) as reporting a specimen of C. fuliginosa from 
Dakto, but Thomas actually referred to it as “‘Crocidura sp.,’’ and we consider 
its identity to be uncertain. Our identifications are based on the discussion of the 
status of C. fuliginosa by Medway (1977), on comparison with specimens (FMNH) 
from China (see Allen 1938), and on data provided by Jenkins (1976). The rec- 
ognition of dracula as a valid species by Lekagul and McNeeley (1977) is based 
on their confusion regarding C. fuliginosa and C. atienuata (see above); our 
examination of specimens supports the recognition by Jenkins (1976) of dracula 
as a subspecies of fuliginosa. 

Specimens examined.—VIETNAM: Con Son Prov.: Con Son Island, | km S, 
0.3 km W Airfield Bldg., elev. 40 m (1 USNM); Annam: Hoi-Xuan (1 FMNH); 
Tonkin: Ba Nam Nhung (1 FMNH); Chapa, elev. 5000 ft (10 FMNH); Lai Chau, 
elev. 500 ft (1 AMNH, 2 FMNH); Muong Mo (1 FMNH). 


Crocidura horsfieldi indochinensis Robinson and Kloss 
Crocidura horsfieldi indochinensis Robinson and Kloss, 1922:88. 


This species is similar in appearance to C. attenuata, but differs in its smaller 
size, proportionately longer tail (tail to head and body ratio averages 0.74), and 
several cranial features noted above (Fig. 2). It is conspicuously different from 
C. fuliginosa in its much smaller size. 

Robinson and Kloss (1922) described indochinensis on the basis of a single 
specimen from Dalat (5000 ft elev.), Langbian Plateau (=Mt. Langbian), Tuyen 
Duc Prov., southern Annam (Vietnam); they provided (among others) the follow- 
ing measurements: tail, 50; hind foot, 12.2; greatest length of skull, 17.2; maxillary 
toothrow to tip of incisors, 7.4; mastoid breadth, 8.1. These measurements fit 
well within the range of those presented here, and it is on this basis that we refer 
our specimens to this taxon. Osgood (1932) reported two specimens from Chapa, 
in extreme northwestern Vietnam, and Anthony (1941) extended the known dis- 
tribution to northeastern Burma. Ellerman and Morrison-Scott (1951) were the 
first to list indochinensis as a subspecies of C. horsfieldi, but offered no expla- 
nation for doing so. This treatment was followed by Jenkins, who stated that the 
holotype of indochinensis was similar to topotypes of C. horsfieldi. We list in- 
dochinensis as a subspecies of C. horsfieldi strictly on that basis. Lekagul and 
McNeely (1977) also followed this treatment, listing specimens from Chengmai 


120 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


and Kao Yai Park in Thailand. Specimens we examined all were taken at or 
above 600 m elevation. We note that C. attenuata and C. h. indochinensis have 
both been taken on Mt. Langbian. 

Specimens examined.—VIETNAM: Tonkin: Chapa, Mt. Fan Si Pan (1 FMNH); 
Binh Dinh Prov.: An Khe Military Base, elev. approx. 600 m (2 USNM); Tuyen 
Duc Prov.: Mt. Langbian, elev. 1700 m (1 USNM). 


Acknowledgments 


We thank G. G. Musser and R. W. Thorington for permission to examine 
specimens under their care, K. F. Koopman and R. S. Voss for their comments 
on the manuscript, Rosanne Miezio for her fine illustrations, and G. Lake for 
assistance with preparing the manuscript. 


Literature Cited 


Allen, G. M. 1938. The mammals of China and Mongolia. Natural History of Central Asia, 11: 
xxv+620 pp.—American Museum of Natural History, New York. 

Anthony, H. E. 1941. Mammals collected by the Vernay-Cutting Burma Expedition.—Field Mu- 
seum of Natural History, Zoological Series 27:37-123. 

Chasen, F.N. 1940. A handlist of Malaysian mammals.—Bulletin of the Raffles Museum, Singapore 
15:xx+ 1-209. 

Ellerman, J. R., and T. C. S. Morrison-Scott. 1951. Checklist of Palearctic and Indian mammals 
1758-1946.—British Museum (Natural History), London, 810 pp. 

Jenkins, P.D. 1976. Variation in Eurasian shrews of the genus Crocidura (Insectivora: Soricidae).— 
Bulletin of the British Museum (Natural History), Zoology 30(7):269-309. 

Lekagul, B., and J. A. McNeely. 1977. Mammals of Thailand.—Kurusapha Ladprao Press, Bang- 
kok. li+758 pp. 

Medway, L. 1977. Mammals of Borneo: field keys and an annotated checklist.—Monographs of the 
Malaysian Branch of the Royal Asiatic Society 7:xii+ 1-172. 

Milne-Edwards, A. 1872. Recherches pour servir a l’histoire naturelle des mammiféres.—G. Mas- 
son, Paris. 

Moore, J. C., and G. H. H. Tate. 1965. A study of the diurnal squirrels, Sciurinae, of the Indian 
and Indochinese subregions.—Fieldiana: Zoology 48:1-351. 

Osgood, W. H. 1932. Mammals of the Kelley-Roosevelts and Delacour Asiatic Expeditions.—Field 
Museum of Natural History, Zoological Series 18(10):193-339. 

Robinson, H. C., and C. B. Kloss. 1922. New Mammals from French Indo-China and Siam.— 
Annals and Magazine of Natural History (9) 9:87-99. 

Thomas, O. 1912. New species of Crocidura and Petaurista from Yunnan.—Annals and Magazine 
of Natural History (8)9:686-688. 

—. 1927. The Delacour exploration of French Indo-China. Mammals.—Proceedings of the 
Zoological Society of London 1927:41-58. 

Van Peenen, P. F. D., P. F. Ryan, and R. H. Light. 1969. Preliminary identification manual for 
mammals of South Vietnam.—United States National Museum, Washington. vit+310 pp. 


(LRH) Museum of Zoology and Division of Biological Sciences, University of 
Michigan, Ann Arbor, Michigan 48109; (RMT) Division of Mammals, Field Mu- 
seum of Natural History, Chicago, Illinois 60605-2496. 


PROC. BIOL. SOC. WASH. 
96(1), 1983, pp. 121-126 


PARVULODESMUS PROLIXOGONUS, A NEW GENUS 
AND SPECIES OF XYSTODESMID MILLIPED 
FROM SOUTH CAROLINA (POLYDESMIDA) 


Rowland M. Shelley 


Abstract.—The minute xystodesmid milliped, Parvulodesmus prolixogonus, is 
described as a new genus and species from a sample collected 30 years ago in 
Abbeville, South Carolina. The principal diagnostic character is a long, slender 
acropodite, which overlaps the sterna of segments 4—6 and curves dorsad into a 
small apical loop. The species is assigned to the tribe Rhysodesmini on the basis 
of gonopodal traits, small body size, and sternal hairs. 


For the past six years I have been investigating the endemic xystodesmid mil- 
liped fauna of piedmont South Carolina, diagnosing three new genera in the tribe 
Apheloriini—Croatania, Brevigonus, and Furcillaria (Shelley 1977, 1980, 1981a), 
and describing other species in a revision of Sigmoria (1981b) and a second paper 
on Brevigonus (1981c). I have visited the area around Abbeville, Abbeville Coun- 
ty, four times searching for a purported new species of the tribe Rhysodesmini 
that Leslie Hubricht collected in the early 1950s. Richard L. Hoffman, who ex- 
amined the milliped at the time of its collection, told me of its existence and that 
it represented a new genus. Unfortunately, no gonopod drawings were available, 
and the preserved specimens were not in his private collection nor in any major 
museum in the eastern United States. Re-collection of the species therefore seemed 
necessary, and I visited Abbeville in spring, summer, and fall, to allow for sea- 
sonal occurrence. These efforts, however, were unsuccessful. 

In 1981 the milliped collection of the late William T. Keeton, former Professor 
of Biology at Cornell University, was transferred to Dr. Hoffman to be incor- 
porated into his holdings. While casually perusing this material one day, I chanced 
upon the long lost sample from Abbeville, which contained two males and two 
females in good condition. However, the gonopods had been removed from one 
male and were not in the vial. Consequently, the critically important genitalic 
features can only be determined from the other male, which I designate the 
holotype. These are the smallest xystodesmids I know of, smaller even than 
Gyalostethus monticolens (Chamberlin) or Pleuroloma pinicola Shelley, and I 
agree completely with Dr. Hoffman that they represent a new genus and species 
in the tribe Rhysodesmini. 


Parvulodesmus, new genus 


Type species.—Parvulodesmus prolixogonus, new species. 

Description.—A genus of minute rhysodesmine xystodesmids with the follow- 
ing characteristics: 

Body composed of head and 20 segments in both sexes; W/L ratio 21-24%. 
Head of normal appearance, smooth; epicranial suture distinct, not bifid; facial 


122 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


setae reduced, epicranial and interantennal absent. Antennae moderately long, 
with 4 small, apical sensory cones. 

Terga smooth, strictures faint. Collum large and broad, but not extending be- 
yond ends of following tergite. Paranota strongly depressed; peritremata flat and 
inconspicuous on all segments, ozopores opening laterad. 

Caudal segments normal for family. 

Pregonopodal sterna of males with variable concave depressions; postgono- 
podal sterna of males and all sterna of females flat and unmodified, with small 
clusters of setae beside leg coxae. Gonapophyses of males short and apically 
expanded. Coxae without ventrodistal spines or tubercles; prefemoral spines short 
and blunt, greatly reduced, absent from some legs. 

Gonopodal aperture relatively large, elliptical. Gonopods with very long ac- 
ropodites, overlapping apically but otherwise parallel to each other, extending 
forward from aperture between legs of segments 6, 5, and 4, lying in sternal 
depressions. Coxae small, without apophyses. Prefemora small, with short, acic- 
ular processes. Acropodites long, thin, and flat, curving dorsad into narrow loops 
apically, with medial subterminal digitiform processes. 

Cyphopodal aperture narrow, encircling second legs. Cyphopods minute but 
with comparatively large, rugulose receptacles overlying ventral valves. Valves 
small and subequal; operculum not detectable but presumed present under free 
end of valves. 

Species.—One is known; others may also occur in piedmont South Carolina. 

Relationships.—Parvulodesmus appears to be most closely related to several 
undiagnosed rhysodesmine species in the Gulf Coastal Plain of southern Alabama. 
These species, which constitute one or possibly two new genera, also have long 
gonopodal acropodites that overlap two or more sterna anterior to the gonopodal 
aperture. 


Parvulodesmus prolixogonus, new species 
Figs. 1-4 


Type specimens.—Male holotype, and one male and two female paratypes (col- 
lection of R. L. Hoffman) collected by Leslie Hubricht, 1 May 1952, from Abbe- 
ville, Abbeville Co., South Carolina. 

Diagnosis.—With the characters of the genus. 

Holotype.—Length 14.7 mm, maximum width 3.1 mm, W/L ratio 21.1%, depth/ 
width ratio 58.1%. Color in life unknown; all specimens completely blanched by 
preservative, without any evidence of stripes or other pigmentation patterns. 

Head capsule smooth, polished; epicranial suture thin but distinct, terminating 
in slight impression in interantennal region, not bifid. Antennae moderately long 
and slender; reaching back to middle of fourth tergite, becoming progressively 
more hirsute distally, with 4 apical sensory cones, no other sensory structures 
apparent, first antennomere subglobose, 2-6 clavate, 7 short and truncate; relative 
lengths of antennomeres 3 > 2 >4>5 =6>1>7. Genae not margined lat- 
erally, with distinct medial impressions, ends broadly rounded and projecting 
slightly beyond adjacent cranial margins. Facial setae as follows: epicranial, in- 
terantennal, frontal, and genal not detected, subantennal 1-1, clypeal about 8-8, 
labral about 12-12. 


VOLUME 96, NUMBER 1 123 


Figs. 1-4. Parvulodesmus prolixogonus. 1, Gonopods in situ, ventral view of holotype. 2, Left 
gonopod of holotype, medial view. 3, Telopodite of the same, lateral view. 4, Distal half of acropodite, 
oblique dorsal view. Scale line for fig. 1 = 1.00 mm; line for other figs. = 1.00 mm for each. 


Terga smooth, polished. Collum broad, ends not produced beyond those of 
following tergite. Paranota strongly depressed, angled sharply ventrad and con- 
tinuing slope of dorsum; anterior corners rounded, caudolateral corners blunt on 
all segments; posterior edges of paranota only slightly discontinuous with dorsum. 
Peritremata thin and inconspicuous, only slightly elevated above paranotal sur- 
face; ozopores located near middle of peritremata, opening laterad. 

Sides of metazonites finely granular, without noticeable grooves or impres- 
sions. Strictures faint, indistinct. Sterna of segments 4-6 with concave central 
depressions, deepest on segment 4, to accommodate stems of gonopodal acro- 
podites; without lobes or other processes. Postgonopodal sterna without lobes, 
caudal edges straight; with shallow transverse grooves between leg pairs and 
short, longitudinal grooves between caudal legs; with 2 to 4 long, slender setae 
in small clusters near coxae of both leg pairs. Pregonopodal legs densely hirsute; 
postgonopodal legs becoming progressively less hirsute caudally. Coxae without 
projections; prefemora apically blunt and rounded, without spines; tarsal claws 
hooked, of normal length on all legs. Hypoproct rounded; paraprocts with margins 
slightly thickened. 


124 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Gonopodal aperture elliptical, with slight indentations anteriolaterad, margins 
elevated above metazonal surface, not thickened. Gonopods in situ (Fig. 1) with 
telopodites projecting ventrad from aperture, extending anteriad in parallel ar- 
rangement over sterna of segments 6, 5, and 4, apices overlapping and interlock- 
ing between third pair of legs. Gonopod structure as follows (Figs. 2-4): Coxa 
small, without apophysis. Prefemur small, with short, acicular process arising on 
dorsal side, extending beyond juncture of prefemur and acropodite for about % 
of length of latter. Acropodite demarcated from prefemur by slight constriction, 
in form of thin, narrow blade extending linearly from prefemur, with apical loop 
of narrow diameter; loop located at about % length, curving dorsad from stem of 
acropodite, with subterminal digitiform projection on medial edge subtending 
small circular area between solenomerite; solenomerite short, apicaily acute, di- 
rected subanteriad. Prostatic groove arising in pit in base of prefemur, crossing 
to dorsal side of acropodite at juncture with prefemur, running along inner surface 
of apical loop and opening terminally on solenomerite. 

Male paratype.—The male paratype agrees with the holotype in all particulars 
except facial setae, which are as follows: epicranial and interantennal absent, 
subantennal 1-1, frontal 1-1, genal 2-2, clypeal about 8-8, labral about 12-12, 
merging with clypeal series and continuing along genal margins for about one- 
third of lengths, about 4 setae on each side. Gonopodal variation is unknown, 
since the paratype gonopods were previously dissected and lost. 

Female paratype.—Length 13.1 mm, maximum width 3.2 mm. W/L ratio 24.4%, 
depth/width ratio 62.5%. Agreeing closely with males in somatic features except 
paranota more strongly depressed, creating appearance of more highly arched 
body, and lacking sternal depressions of pregonopodal segments of males. Cy- 
phopods with comparatively large, subtriangular receptacle visible in aperture, 
lying over and obscuring valves, surface rugulose; valves small and subequal, 
located dorsad to receptacle, opening anteriad, surfaces finely granulate. 

Distribution.—Known only from the type locality. 

Remarks.—The prefemoral process of P. prolixogonus can be easily over- 
looked or mistaken for a seta or cotton fiber, when the gonopod is placed on 
cotton for drawing. It is colorless, translucent, and closely appressed to the pre- 
femur and base of the acropodite. Its shortness relative to the extremely long 
acropodite makes it look like a macroseta, and it blends into the acropodite and 
becomes nearly invisible in any view except medial, which reveals the narrow 
gap between them. The process is also very fragile and can be easily broken 
during gonopod dissection. 

Parvulodesmus prolixogonus is assigned to the tribe Rhysodesmini because of 
the small body size, the hairs on the sterna, the parallel arrangement of the 
gonopods, and the acicular prefemoral process. These traits are all more typical 
of the Rhysodesmini than any other southeastern xystodesmid tribe. Small body 
size is shared with the genera Gyalostethus, Caralinda, and the undescribed 
Alabama taxa; hirsute sterna are exhibited by two species of Pleuroloma; and all 
rhysodesmine genera except Caralinda display acicular prefemoral processes and 
parallel gonopods. One rhysodesmine trait not observed was the sternal remnant 
connecting the gonopodal coxae. I neglected to check this character before dis- 
secting the gonopods of the holotype, and the nature of the remnant will have to 
be determined from fresh material. 


VOLUME 96, NUMBER 1 125 


In trying to discover more specimens near Abbeville, I checked a wide variety 
of biotopes in every habitat I could think of. I also wrote the collector, but he 
could only remember that the sample was taken near the railroad. Many of the 
places I investigated were urbanized sites in or near Abbeville, and one trip came 
during the last week of April, the same time of year that the type sample was 
collected. Thus, I think I checked, at the proper time of year, the kinds of sites 
Hubricht did. On the thought that Abbeville might be at the range periphery where 
the species is least common, I investigated distant parts of the county and adja- 
cent McCormick, Greenwood, and Anderson counties, to try to find an area of 
greater abundance. Although not specifically looking for Parvulodesmus, I have 
also investigated thoroughly during the past six years the entire Piedmont of South 
Carolina and, to a lesser extent, the tier of counties along the southern side of 
the Savannah River in Georgia. However, trying to find such a tiny animal in 
such a large area with no habitat information is exceedingly difficult, and it is not 
surprising that none of these activities were successful. I think that the situation 
with Parvulodesmus may be similar to that of Caralinda in southern Georgia, 
Alabama, and neighboring Florida. In this genus single species were described 
by Hoffman (1978) and me (1979), but I have recently discovered a large amount 
of Caralinda from the Florida panhandle in the Florida State Collection of Ar- 
thropods, containing two or three more new species. Thus, Caralinda consists 
of four to five species and occupies a much larger area than previously thought. 
Similarly, Parvulodesmus may also consist of two or more species and occupy a 
much larger area in South Carolina than just Abbeville County. However, I have 
exhausted all my knowledge and field experience trying to find more individuals, 
and my only further thought is that May might be the time of least abundance. 
Perhaps the species of Parvulodesmus, like Nannaria conservata Chamberlin 
(Shelley 1975), are most abundant during the winter months and are prevalent in 
December, January, and February. I leave this idea for future investigators, who 
may be challenged by the rarity of Parvulodesmus, or local biologists, who live 
in the Abbeville-Greenwood area and can sample in winter and continuously 
throughout the year. A continuous, long-term effort of this kind may be required 
to find Parvulodesmus again. 


Acknowledgments 


I thank Richard L. Hoffman, for the privilege of describing these new taxa, 
and Renaldo G. Kuhler, N.C. State Museum scientific illustrator, who prepared 
Figure 1. This study was aided by financial assistance by the National Science 
Foundation, Grant No. DEB 7702596. 


Literature Cited 


Hoffman, Richard L. 1978. A new genus and species of rhysodesmine milliped from southern 
Georgia (Polydesmida: Xystodesmidae).—Proceedings of the Biological Society of Washington 
91:365-373. 

Shelley, Rowland M. 1975. The identity of Nannaria conservata Chamberlin, with notes on an 
abnormal male and descriptions of two new species of Nannaria from North Carolina (Diplo- 
poda: Polydesmida: Xystodesmidae).—Proceedings of the Biological Society of Washington 
88:179-188. 


126 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


———. 1977. The milliped genus Croatania (Polydesmida: Xystodesmidae).—Proceedings of the 
Biological Society of Washington 90:302-325. 4 

——.. 1979. A new milliped of the genus Caralinda from north Florida (Polydesmida: Xystodes- 
midae).—Florida Entomologist 62:184—187. 

———. 1980. The status of Cleptoria shelfordi Loomis, with the proposal of a new genus in the 
milliped family Xystodesmidae (Polydesmida).—Bnmleyana 3:31—42. 

——. 198la. A new xystodesmid milliped genus and three new species from piedmont South 
Carolina (Polydesmida).—Proceedings of the Biological Society of Washington 94:949_967. 

—. 198lb. Revision of the milliped genus Sigmoria (Polydesmida: Xystodesmidae).—Memoirs 
of the American Entomological Society No. 33, 140 pp. 

—.. 198lc. A new milliped of the genus Brevigonus from South Carolina, with comments on 
the genus and B. shelfordi (Loomis) (Polydesmida: Xystodesmidae).—Brimleyana 6:51—60. 


North Carolina State Museum of Natural History, P.O. Box 27647, Raleigh, 
North Carolina 27611. 


PROC. BIOL. SOC. WASH. 
96(1), 1983, pp. 127-133 


CURIDIA DEBROGANIA, A NEW GENUS AND SPECIES 
OF AMPHIPOD (CRUSTACEA: OCHLESIDAE) 
FROM THE BARRIER REEFS OF BELIZE, 
CENTRAL AMERICA 


James Darwin Thomas 


Abstract.—The family Ochlesidae is amended to include the new genus Curi- 
dia, which differs from all other members by possessing maxillipedal palps. Cur- 
idia debrogania is described and compared to other genera and species within 
the family Ochlesidae. 

Curidia debrogania is plesiomorphic by possession of maxillipedal palps, sug- 
gesting this member of the small, cryptic family might have originated in the 
tropical Western Atlantic. Distribution records and ecological notes are included. 


Family Ochlesidae 


Accessory flagellum absent; palp of maxilliped absent or uniarticulate; mouth- 
parts projecting subconically; mandible with molar reduced or lacking; coxae I- 
4 subacuminate; telson entire. 


Curidia, new genus 


Diagnosis.—Article 1 and 2 of antenna | each with long ventrodistal cusp, 
peduncular article | longer than 2, flagellum 4-articulate, article 2 longest, ex- 
ceeding peduncular article | in length. Coxa | rounded ventrally, coxae 2 and 3 
dominant, elongate. Upper and lower lips apically acute; mandible with molar; 
first maxilla with palp, outer plate styliform; second maxilla with both plates 
styliform, outer plate longest; maxilliped with elongate inner plate, outer plate 
slightly armed, palp present, consisting of slender unarmed article bearing ter- 
minal seta. 

Etymology.—The name Curidia is from the Greek, Kouridios, ““wedded,”’ 
(gender feminine), and is in reference to Stebbing’s original designation of the 
type-genus Ochlesis, which means “‘disturbance.’’ This designation referred to 
the lack of maxillipedal palps in the type species, O. innocens Stebbing, 1910. 
The presence or absence of these palps is a primary distinction between the 
hyperiidean and gammaridean amphipods. Hence, Stebbing felt that the lack of 
these structures was a significant “‘disturbance,” or ‘‘disruption’’ among the 
Gammarideans. The presence of maxilliped palps in the genus Curidia allows the 
family Ochlesidae to be included without exception among the Gammarideans. 


Curidia debrogania, new species 


Description.—Male: body strongly compressed and very thin dorsally; pleonite 
2 with rounded process dorsally; urosomite | thin and elongate; epimeron 3, 
posteroventral margin with upturned hook; head small with acute lateral cephalic 
lobe, eye prominent, composed of 21 ommatidia; article 1 of antenna | longer 


128 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. |. Curidia debrogania, holotype, male, 1.59 mm, JDT Bel 48, Carrie Bow Cay, Belize. 


than article 2, articles | and 2 each with large ventrodistal cusp, cusp of article 
1 extending to end of article 3, cusp of article 2 extending halfway along flagellum; 
accessory flagellum absent; primary flagellum 4-articulate, article 2 elongate, api- 
cal and ventral margins with long aesthetascs; antenna 2 shorter and less robust 
than antenna 1, article 4 with ventral cusp extending halfway along article 5; 
flagellum 4-articulate, article 2 slightly smaller than article 1; upper lip large, 
triangular; mandibles massive, styliform, incisors simple, left mandible with small 
lacinia mobilis, molar small, circular, weakly triturative, ratio of palp articles 1- 
3, 35:45:57, article 3 with double row of small facial ridges and 3 short spines 
distally; lower lip slender, apices extended, mandibular projections produced and 
narrowly rounded; maxilla |, inner plate small with | apical and | subapicomedial 
seta, outer plate attenuated, styliform and slightly recurved distally, medial mar- 
gin bearing 5 hooked spines subapically, preceded proximally by marginal row 
of fine setae, palp uniarticulate, borne on raised process and bearing single long 
terminal seta; maxilla 2, inner plate styliform with 7 pectinate spines (5 medial, 
1 apical, 1 subapical) and 4 short setae on medial margin, outer plate longer and 
thinner than inner plate, bearing several long apical setae, plus single basofacial 
spine and 6 short setae on lateral margin; maxilliped, inner plate thin, elongate 
(0.77 times outer plate), with 3 facial and 2 distomedial plumose setae, short apical 
nail, and numerous short setae on distolateral margin, outer plate curled orally, 
apical margin rounded with 4 subapical spines and plumose seta, 2 mediofacial 
setae on oral margin, palp uniarticulate, bearing long terminal seta extending 
beyond outer plate; coxa | small, dentate on anteroventral margin; coxa 2 elon- 
gate, anterior margin produced; coxa 3 also elongate with straight anterior margin, 
posterior margin produced; coxa 4 smaller than coxae 1-3, anteroventral margin 
truncate; coxae 5 and 6 similar, with posteroventral margins produced; coxa 7 


129 


VOLUME 96, NUMBER 1 


W) 
uf COTTA 


iy, 


ty 
1 
‘1, 

"1 f1, 


"Hin 
Vay 
u 


K 


Curidia debrogania, A-I, holotype, male, 1.59 mm, JDT Bel 48: A, Maxilla 1; B, Maxilla 


Fig. 2. 
2; C, Maxilliped outer plate; D, Lower lip; E, Maxilliped; F, Maxilliped inner plate; G, Right mandible; 


H, Mandibular Palp; I, Lower lip. J-K, paratype, female, 1.40 mm: J, Antenna 1; K, Antenna 2. 


130 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


= 


Fig. 3. 


Curidia debrogania, holotype, male, 1.59 mm, JDT Bel 48: A, Gnathopod 2; B, C, D, 
Uropods 3, 2, 1; E, Gnathopod 1; F, Gnathopod 2; G, Telson; H, Gnathopod 1. 


with slightly rounded posterior concavity; gnathopod | simple, article 2 slightly 
inflated, articles 3 and 4 subequal and elongate, article 5 longer than 6, latter with 
2 plumose setae and 2 spines near joint, dactyl with accessory process and 2 long 
plumose setae; gnathopod 2 basis slightly inflated, articles 3 and 4 short, subequal, 


VOLUME 96, NUMBER 1 131 


article 4 extending distoventrally, article 5 elongate and distoventrally produced, 
article 6 elongate, shorter than 5, dactyl bifid with accessory process; pereopods 
3 and 4, article 4 with anterodistal process extending approximately halfway along 
article 5, article 6 with several posterior spines, dactyl massive, elongate; per- 
eopods 5-7 similar in structure to pereopods 3 and 4; pleonite 2 with obtuse 
middorsal hump, pleonite 3, lateral margins with strong upturned tooth; urosomite 
1 elongate; uropod | reaching beyond uropods 2 and 3, peduncle elongate, sub- 
equal to rami, with single apicolateral spine and 3 medial marginal spines, rami 
lanceolate, subequal, outer ramus with 4 lateral marginal and 2 medial marginal 
spines, inner ramus with paired medial and lateral marginal spines, both margins 
of outer ramus and medial margin of inner ramus finely serrate; uropod 2 peduncle 
with | apicolateral, | midlateral, and | subapicomedial spine, outer ramus sub- 
equal to peduncle, with 2 spines on lateral margin and single distomedial spine, 
inner ramus 1.36 times the outer, paired spines on distolateral and distomedial 
margins, all margins of both rami finely serrate; uropod 3 peduncle with short 
subapicolateral spine, outer ramus short, with a single spine on lateral margin, 
inner ramus 1.43 times the outer with 2 lateral and 2 medial marginal spines, 
entire lateral margin of outer ramus and both margins of inner ramus finely ser- 
rate; telson entire, elongate, with 2 plumose setae. 

Female.—Apart from the presence of brood plates, the only apparent differ- 
ences are in antenna |. Females of C. debrogania have smaller cusps and lack 
ventral setae on the flagellum, all setae being apical. Ovigerous specimens have 
been taken from Biscayne Bay, Florida, and Belize as small as 1.40 mm. 

Etymology.—Curidia debrogania is named in honor of Debra L. Rogan, Sep- 
tember 8, 1950 to December 25, 1981. 

Types.—Deposited in the United States National Museum of Natural History 
collections, holotype, USNM 191041, male, 1.59 mm; paratype, USNM 191042, 
female, 1.40 mm. 

Type-locality.—JDT Belize 48, Carrie Bow Cay, Belize, 7 June 1980, 6 m, partly 
cemented coral rubble in patch reef area just inside channel between Carrie Bow 
Cay and Water Cay, 16°48’N, 80°05’W. 

Color.—White laterally, blending to translucent brown along dorsal and ventral 
margins of body. Pereonites 1-7, antenna 2, gnathopods, and pereopods 3-7 with 
distinct purple or deep maroon banding. 

Distribution!—Barrier reefs of Belize, to Biscayne Bay, Florida, sublittoral, 2- 
20 m, usually associated with macroalgae. 

Ecology.—Curidia debrogania is usually associated with coral reef habitats 
where it is found in areas of high current velocity or wave surge (i.e., lagoonal 
channels, or in the forereef areas). The mouthparts suggest a parasitic existence 
although no documentation exists to support this thesis at present. The massive 
dactyls on pereopods 3-7 could be used to hold on to a host in areas of high 
current velocities, or may aid in forcing the conical mouthparts into host tissues. 
The distinctive color pattern suggests that C. debrogania may be a mimic of some 
species of micromollusc. Specimens of C. debrogania are usually taken from 
rocky outcrops that have attached macroalgae. 

Discussion.—While C. debrogania is unique among ochlesids in possessing 
maxilliped palps, it appears, in other external characters, closest to Ochlesis alii 
Barnard, 1970, from Hawaiian waters, and O. innocens Stebbing from the Aus- 


132 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


tralian littoral. Curidia debrogania, O. alii, and O. innocens are the only ochlesids 
having both antennal cusps and teeth on the posterior margin of epimeron 3. 
Males of O. alii (not at present described) should prove to have somewhat more 
prominent antennal cusps than females and therefore approach the condition of 
C. debrogania males. Curidia debrogania differs from O. alii in 1) the longer 
cusps on antennae | and 2; 2) the more robust palp of maxilla 1; 3) less spinose 
and laceolate uropods. Ochlesis innocens resembles C. debrogania in the large 
antennal cusps, but differs in 4) shorter peduncular article 2 and much longer 
flagellar article 2 of antenna |; 5) absence of a palp on maxilla | (if Pirlot’s 
identification of O. innocens from Isles Aru is correct, then a rudiment of a palp 
is present); 6) absence of a cusp on antenna 2; 7) ventral margin of coxa | acute 
versus rounded in C. debrogania. 

Two of the four remaining species of ochlesids, O. levetzowi Schellenberg, 
1953, and O. eridunda Barnard, 1972, lack antennal cusps and a tooth on epi- 
meron 3, and in general body appearance are not as laterally flattened and dorsally 
acute as those species with antennal cusps and a tooth (or teeth) on epimeron 3. 
Oclesis lenticulosa K. H. Barnard, 1940, from the littoral of South Africa lacks 
antennal cusps, but has a tooth on epimeron 3. Oclesis meraldi Barnard, 1972, 
from the Australian littoral differs so grossly in external morphology and orna- 
mentation from all other ochlesids that it will be used as the type-species for a 
new genus (Barnard, pers. comm.). 

The presence of maxillipedal palps in C. debrogania suggests it to be more 
primitive (plesiomorphic) than other members of the family, all of which lack this 
character. This would suggest a Western Atlantic origin for the ochlesids, whereas 
Pacific and Indo-Pacific forms would be of more recent origin in having lost the 
maxillipedal palps. This hypothesis will remain unproven until additional material 
can be studied from wide geographical areas. Curidia debrogania is also the 
smallest representative in the Ochlesidae, with fully ovigerous females being tak- 
en at a length of 1.40 mm. Ochlesids have been overlooked by most investigators 
because of their minute size (most species) and cryptic habitat. Not until thorough 
sampling of all tropical reef areas is conducted will the zoogeograpical affinities 
and ecology of ochlesids be better understood. 


Acknowledgments 


I wish to thank Mike Carpenter of the Smithsonian Institution for tireless as- 
sistance in the field under trying conditions; Dr. J. L. Barnard of the Smithsonian 
Institution for critically reviewing the manuscript; the Smithsonian’s Investiga- 
tions of Marine Shallow Water Ecosystems (IMSWE), which is partly funded by 
a grant from the Exxon Corporation, for supporting the author at its field labo- 
ratory at Carrie Bow Cay, Belize. This research was funded by grants DEB 
7920534 and DEB 8121128 from the National Science Foundation. This paper 
forms IMSWE contribution number 120. 


Literature Cited 


Barnard, J. L. 1970. Sublittoral Gammaridea (Amphipoda) of the Hawaiian Islands.—Smithsonian 
Contributions to Zoology 34:1—286, 180 figs. 


VOLUME 96, NUMBER 1 133 


———. 1972. Gammaridean Amphipoda from Australia, Part I.—Smithsonian Contributions to 
Zoology 103:1—333, 194 figs. 

Barnard, K. H. 1940. Contributions to the crustacean fauna of South Africa, XII: Further additions 
to the Tanaidacea, Isopoda, and Amphipoda, together with keys for the identification of hith- 
erto recorded marine and freshwater species.—Annals of the South African Museum 32:38 1- 
543, 35 figs. 

Schellenberg, A. 1953. Erganzungen zur Amphipoden fauna Stidwest-Afrikas nebst Bemerkungen 
uber Bratraumbildung.—Mittheilungen aus dem Zoologisch Museum in Berlin 29:107—126, 7 
figs. 

Stebbing, T. R. R. 1910. Crustacea. Part 5. Amphipoda. Jn Scientific Results of the Trawling 
Expedition H.M.C.S. Thetis —Memoirs of the Australian Museum 4(2):565—658, pl. 47-60. 


Newfound Harbor Marine Institute, Big Pine Key, Florida 33043. 


PROC. BIOL. SOC. WASH. 
96(1), 1983, pp. 134-144 


PROECHIMYS SEMISPINOSUS (RODENTIA: 
ECHIMYIDAE): DISTRIBUTION, TYPE 
LOCALITY, AND TAXONOMIC HISTORY 


Alfred L. Gardner 


Abstract.—The type locality of Proechimys semispinosus (Tomes) has been 
alleged to be Gualaquiza, in the lowland forests of southeastern Ecuador. A 
review of the travels of the collector Louis Fraser, and examination of the content 
of reports based on the mammals he collected, plus the comparison of the type 
specimen with samples representing several species of Proechimys from Central 
and South America, indicate that the type did not come from Gualaquiza. The 
type locality of P. semispinosus is corrected to Esmeraldas, on the Pacific coast 
of Ecuador. 


The specimens on which Tomes (1860b) based the name Echimys semispinosus 
were collected by Louis Fraser in Ecuador and alleged (Allen 1916) to have come 
from Gualaquiza, a community on the Rio Santiago in the Province of Morona- 
Santiago. This site (03°24’S, 78°33’W fide Paynter and Traylor 1977) is east of 
the Andes, at 750 meters elevation, in the tropical zone of the western Amazonian 
forests. I have concluded, however, from my examination of the skull of the type 
(British Museum [Nat. Hist.] 7.1.1.173), that it is unlike skulls of any of the 
species of Proechimys known to me from South America east of the Andes. 
Instead, it has the cranial features that characterize populations of Proechimys 
found west of the Andes along the Pacific lowlands of Colombia and Ecuador 
and northward through Central America to Honduras that currently are known 
in the literature as either Proechimys centralis or P. semispinosus. 

The present report is the result of my examination of specimens and search of 
the literature in an attempt to determine the provenance of the specimens that 
Tomes used in describing P. semispinosus. In addition to the examination and 
comparison of specimens, including types, I have explored three main sources 
of information: the published history of the use of the name P. semispinosus, 
accounts of Fraser’s travels in Ecuador, and the content of reports on the mam- 
mals Fraser collected. 


Taxonomic History 


1860 
Tomes described Echimys semispinosus based primarily on one of “‘three spec- 
imens . . . received in spirits, all of which were females; one of them contained 


two young ’’ (1860b:267—268), and figured the skull of one of these. The specimens 
had been collected by Fraser at an unspecified locality in Ecuador. Tomes intro- 
duced this account by stating (p. 265), “‘In my first notes [Tomes 1859] on Mam- 
mals, collected by Mr. Fraser, I included the Echimys cayennensis from the 
examination of a specimen which had lost the tail, and was otherwise in an 
unsatisfactory condition. Other and better specimens of Echimys having been 


VOLUME 96, NUMBER 1 135 


received, I have been able to make out clearly that they represent a new and 
well-marked species, and that the former specimen was similar to them.”’ 


1889 
True recorded Echinomys semispinosus from Nicaragua and Costa Rica. 


1896 

Thomas described Echinomys centralis (type locality: San Emilio, south end of 
Lake Nicaragua) and said that True’s (1889) report was based on the same species. 
Thomas also stated (p. 312), “Both in the Merida and Bogota collections there 
occur specimens of the genus Echinomys which, on account of their (in compar- 
ison with other species) ‘very dark brown colour’ and other characters, may fairly 
be assigned to E. semispinosus, Tomes, especially if, as seems to be the case, 
Tomes’s woodcut of the skull is incorrect in details.’’ Thus Thomas assigned 
seven specimens (p. 313) from Colombia to E. semispinosus. 


1897 

Thomas, in his description of Echimys gymnurus (=Hoplomys gymnurus), re- 
marked (p. 551), ‘‘This handsome Echimys differs from Tomes’s E. semispinosus, 
also from Ecuador, but without exact locality, by its richer rufous colour... . 
In all these points the Bogota Echimys assigned last year to E. semispinosus 
agrees much better with Tomes’s description than does that [E. gymnurus] brought 
home by Mr. Rosenberg, in spite of the Ecuadoran locality of the latter.’’ 


1898 

Thomas recanted his assignment of the Bogota specimens (which he here named 
Echimys chrysaeolus) to E. semispinosus Tomes, and decided that E. semispi- 
nosus was allied with the Nicaraguan specimens he had already described (1896) 
as E. centralis. This resulted from the discovery of *‘a dark brown Echimys 
marked, in what appears to be a hand writing similar to that on some of Fraser’s 
labels, “Echimys semispinosus,’ >’ among specimens that had been overlooked for 
many years in rooms of the Zoological Society. Thomas stated (p. 244), ‘This 
[specimen] so precisely agrees with Tomes’s description of that animal, that there 
can be no reasonable doubt that it is one of the original specimens collected by 
Fraser and referred to in Tomes’s paper.’’ The label may have lacked notation 
of locality for Thomas failed to mention one. 


1899 
Allen proposed the name Proechimys (type-species, Echimys trinitatus Allen and 
Chapman) to include, among others, the species P. semispinosus. 


1900 

Thomas described Proechimys rosa (type locality, Santa Rosa, Prov. El Oro, 
Ecuador), P. centralis panamensis (type locality, ‘‘Savanna near Panama’’) and 
P. centralis chiriquinus (type locality, Bugava [=Bugaba], Prov. Chiriqui, Pan- 
ama). Thomas allied P. semispinosus with P. decumanus (described by Thomas 
1899, from Chongon, Prov. Guayas, Ecuador), but said that P. rosa was allied 
to the Central American species P. centralis. 


1901 
Bangs described Proechimys burrus (type locality, Isla San Miguel, Golfo de 
Panama), which he allied with P. centralis. 


136 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


1905 
Bangs described Proechimys gorgonae (type locality, Isla Gorgona Colombia), 
which he allied with P. centralis panamensis. 


1911 

Thomas described P. semispinosus calidior (type locality, San Javier, Prov. Es- 
meraldas, Ecuador), calling it ‘‘A rufous lowland representative of true semi- 
spinosus.’ In a footnote (p. 254), he said that the type of P. semispinosus was 
British Museum (NH) number 7.1.1.173, which means that the specimen was the 
173rd cataloged on | January 1907. 


1914 

Hollister described Proechimys rubellus from Costa Rica. He said the type came 
from the Angostura Valley, but as True (1889:467) had stated earlier and as 
indicated on the label of the holotype, the actual collection site is Pacuare. 


1914 

Thomas described P. centralis colombianus (type locality, Condoto, Depto. Cho- 
co, Colombia) and compared it to ‘“‘Proechimys xanthaeolus,’’ apparently a lap- 
sus for P. chrysaeolus. 


1916 

Allen wrote (pp. 206-207), ‘“‘The type locality of Echimys semispinosus Tomes, 
according to Thomas (on the back of label), is Gualaquiza, Ecuador, this being 
the specimen from which the skull was figured. Three specimens of this species 
were received ‘in spirits, all of which were females; one of them contained two 
young’ (Tomes, P. Z. S., 1860, p. 267). Two of these specimens are now in the 
British Museum, ‘evidently skinned out of spirit and much discolored,’ as stated 
on the labels. I omitted to examine the type skull figured by Tomes, but from an 
examination of Tomes’s figure it is too adult to have belonged to either of these 
two skins, which are only about half grown. The skin of the third specimen, 
doubtless the adult female mentioned by Tomes, and the one to which the figured 
skull belonged, has apparently been lost.’ 


1920 

Goldman arranged all of the named forms of Central American Proechimys as 
subspecies of P. semispinosus, except for P. centralis chiriquinus, which he 
treated as a synonym of P. semispinosus panamensis. 


1935 
Tate in his review of the taxonomy of Proechimys, listed P. semispinosus with 
western Amazonian species on the basis of the alleged type locality. 


1937 
Bole described P. semispinosus goldmani from Altos Cacao, Prov. Veraguas, 
Panama. 


1940 

Ellerman listed all named forms of Proechimys semispinosus as subspecies or 
synonyms of P. cayennensis (Desmarest), but remarked that if this be incorrect, 
burrus, centralis, panamensis (with chiriquinus a synonym), rubellus, colombi- 
anus, and calidior should be considered races of P. semispinosus. 


VOLUME 96, NUMBER | 137 


1944 

Osgood said that the type of Proechimys semispinosus was a skull only and that, 
as the type locality was in southeastern Ecuador, populations from northern Peru 
(Huallaga River) probably were identical or closely related. He also assigned a 
series of specimens from Lagunas in east central Peru to P. semispinosus. Osgood 
believed P. semispinosus calidior to be a “‘slight subspecies of semispinosus”’ 
allied with P. rosa and Central American forms. 


1946 
Kellogg described P. semispinosus ignotus from Isla San José, Golfo de Panama, 
Panama. 


1946 
Goodwin treated Costa Rican centralis, panamensis, and rubellus as subspecies 
of P. cayennensis. 


1948 

Hershkovitz considered P. semispinosus a subspecies of P. guyannensis (E. 
Geoffroy), the senior synonym of P. cayennensis (Desmarest). He described (p. 
138) P. quadruplicatus from Isla Llunchi, Rio Napo, Ecuador, and referred to it 
the specimens from Lagunas, Peru, that Osgood (1944) had assumed to be rep- 
resentative of P. semispinosus. Based on the enamel patterns of the cheekteeth, 
he said that two kinds of Proechimys occur in Central America; those assignable 
to P. guyannensis as subspecies, and others (including ignotus, which he elevated 
to species rank) referable to the P. quadruplicatus group. 


1948 

Moojen treated P. semispinosus as a species to which he assigned all of the named 
forms of Proechimys from Central America and western Colombia and Ecuador 
as well as the Amazon basin populations he believed were represented by the 
names gularis, hilda, kermiti, liminalis, and amphichoricus. Moojen described 
the last two taxa as new subspecies. 


52 

Hall and Kelson followed Moojen (1948) and recognized burrus, centralis, gold- 
mani, ignotus, panamensis (chiriquinus a synonym), and rubellus as subspecies 
of P. semispinosus. 


1961 

Cabrera followed Moojen’s (1948) application of P. semispinosus to the assign- 
ment of names to South American Proechimys, with the following exceptions: 
colombianus, decumanus, and gorgonae treated as subspecies of P. guyannensis; 
gularis (sensu Moojen 1948; not of Thomas 1911) assigned to P. quadruplicatus 
Hershkovitz. 


1966 
Handley considered the taxa burrus, goldmani, ignotus, and panamensis to be 
subspecies of Panamanian P. semispinosus. 


1972 
Patton and Gardner used the name P. semispinosus for Costa Rican topotypes 
of P. rubellus. 


138 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


1976 
Handley identified some Venezuelan populations as P. semispinosus. 


1976 

Reig and Useche used P. centralis for the Costa Rican Proechimys that Patton 
and Gardner (1972) had called P. semispinosus, stating that they preferred to 
follow Thomas’s original application of P. centralis to all Central American mem- 
bers of the genus. 


1981 
Hall applied P. semispinosus in the same way the name was used by Hall and 
Kelson (1959). 


Fraser’s Travels in Ecuador 


I have gleaned information on Louis Fraser’s travels in Ecuador, from the time 
he arrived in Guayaquil, 20 September 1857, until he left Esmeraldas at the end 
of December 1859, from three reports by Tomes (1859, 1860a, b) on the mammals, 
several reports on the birds by Sclater (1859a, b, c, 1860a, b, c, d, e), and pub- 
lished extracts of letters written from Ecuador by Fraser (1858a, b, 1859a, b, c, 
d, 1860). 

After a brief stay in Guayaquil, Fraser proceeded to Cuenca, arriving on 6 
October 1857 and remaining there through November. Then he journeyed to 
Gualaquiza where he worked from December 1857 through February 1858, except 
for a two-week period in Zamora during January. 

Fraser left Gualaquiza for Cuenca on | March (arriving 5 March) where he 
collected through April and May before proceeding to Riobamba. He worked in 
or near Riobamba during June and again in August on his return from Quito 
before continuing on to Pallatanga. His collections in the Pallatanga area, with 
trips to Chillanes, were made from the latter part of August through December 
1858. 

Leaving Pallatanga in mid-January 1859, Fraser collected on the slopes of 
Chimborazo north of Riobamba in February on his way to Quito. March, April, 
May, and part of June were spent in the vicinity of Quito and on the slopes of 
Pichincha as well as on the western slopes of the Andes northwest of Quito. 

Fraser traveled southward from Quito to Babahoya where he stayed from 10 
July through part of September. Then he went on to Guayaquil where, in the 
beginning of October, he left by ship for Esmeraldas. Fraser remained in Es- 
meraldas until the end of December and then left Ecuador for Guatemala. 


Tomes’ Reports on Fraser’s Ecuadoran Mammals 


Like many naturalist travellers of his day, Fraser collected a variety of verte- 
brates, invertebrates, and plants, but concentrated on birds. Among the several 
reports based wholly or in part on the mammals he collected in Ecuador are three 
major ones by Tomes published in the Proceedings of the Zoological Society of 
London (1859, 1860a, b). 

In the first report, ‘‘Notes on a Collection of Mammalia made by Mr. Fraser 
at Gualaquiza,’’ Tomes (1859) listed 17 taxa (some unidentified). Among these 
was (p. 548) “‘Echimys cayenensis, Geoff. A thickly-spined and rather large ex- 


VOLUME 96, NUMBER 1 139 


ample, apparently somewhat more strongly tinged with rufous than the illustration 
given by Mr. Waterhouse, or than either of those given by M. Pictet.’’ The list 
was incomplete, however, for Fraser (1858b:6158) said in a letter dated Guala- 
quiza, 13 February 1858, that he had ‘“‘skulls of tapir and white-lipped peccary,”’ 
which are missing from Tomes’ list. Moreover, not all of the animals that Tomes 
listed actually came from Gualaquiza. Fraser (1858a:5942) stated in a letter from 
Cuenca dated 21 October 1857 that he had ‘‘three or four Mammalia’ on hand 
before he went into the southeastern lowlands of Ecuador. While at Gualaquiza, 
Fraser took a side trip to Zamora and collected mammals there. Nevertheless, 
there is no reason to question Gualaquiza as the place of origin of the rat Tomes 
(1859:548) reported as ““Echimys cayenensis.”’ 

The next report (Tomes 1860a), ‘“‘Notes on a Second Collection of Mammalia 
made by Mr. Fraser in the Republic of Ecuador,” listed 15 identified and un- 
identified species, two of which (Hesperomys latimanus and H. minutus) were 
described as new. At the end of the report, Tomes described two species (Hes- 
peromys bicolor and H. aureus) based on specimens previously listed in his first 
report (Tomes 1859). With the exception of these two last named species and the 
Diphylla ecaudata (which came from *‘Rio Napo”’ and had been given to Fraser), 
the mammals of the second report were believed by Tomes (1860:211), “‘to have 
been collected at Pallatanga on the western slope of the Cordillera: but the exact 
locality is not certain, from the specimens having been unfortunately mixed to- 
gether.’’ Tomes gave evidence that Pallatanga was the origin of some of this 
material. Some specimens, however, may have come from Chillanes (known to 
have been visited by Fraser during his stay at Pallatanga), or from the vicinity of 
Riobamba where Fraser worked before going to Pallatanga (but there is no record 
that he collected mammals there). 

The final report (Tomes 1860b), ‘““Notes on a Third Collection of Mammalia 
made by Mr. Fraser in the Republic of Ecuador,’’ contained 21 taxa, of which 
Hesperomys caliginosus, H. albigularis, and Echimys semispinosus were de- 
scribed as new. Pallatanga was indicated as the source of H. albigularis, but no 
locality data were given for the other two newly described species. Actually, 
localities were mentioned for only nine of the 21 taxa that Tomes listed. However, 
attention to dates, information from letters, and subsequent designations of type 
localities result in coastal Ecuador indicated for 1; Cuenca for 2; Gualaquiza, 3; 
Zamora, 2; Pallatanga, 2; Esmeraldas, 5; and unknown, 6. Of the three taxa from 
Gualaquiza, two are represented by the tapir and white-lipped peccary skulls first 
mentioned in Fraser’s (1858b:6158) letter from Gualaquiza. The third is Echimys 
semispinosus represented by three specimens including the type, which Allen 
(1916) said, based on Thomas’ determination, came from Gualaquiza. 


Discussion 


Fraser collected four specimens of Proechimys: the specimen from Gualaquiza 
first reported by Tomes in 1859 as Echimys cayenensis and three females in spirits 
reported without locality by Tomes in 1860. One of these three, an adult with 
two embryos, was the subject of the illustrations and measurements given by 
Tomes (1860b) in his description of Echimys semispinosus and to which he said 
the Gualaquiza specimen was ‘‘similar.’’ It is apparently this specimen that was 


140 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


later given British Museum (NH) catalog number 7.1.1.173, and is now labelled 
as the type of Echimys semispinosus Tomes. 

The skin of the holotype now lacks a tail. The tail was present when Tomes 
(1860b) described P. semispinosus because he said it measured 5 inches, 6 lines 
(=139.7 mm). The skull of the holotype is in good condition. These clues confirm 
that the skin and skull presently labelled as the holotype of P. semispinosus do 
not represent the animal Tomes reported in 1859 from Gualaquiza, because Tomes 
(1860b) commented on the unsatisfactory condition of that specimen and noted 
that it lacked a tail. 

It can be inferred from Thomas’ (1896, 1897) writings that the holotype of P. 
semispinosus had been lost for some time before it was located among some 
specimens in rooms of the Zoological Society of London (Thomas 1898). Prior 
to its discovery, Thomas had assumed that material he later named P. chrysaeolus 
(Thomas, 1898) was representative of P. semispinosus. His confidence in that 
assumption was sufficiently strong for him to suggest that certain details in Tomes’ 
figure of the skull were incorrect (Thomas 1896:312). I presume that the details 
Thomas was referring to included the strongly-developed parietal ridges. A single, 
strongly-developed ridge across the parietals is a feature characteristic of all 
populations of Proechimys found in Central America and in northern South Amer- 
ica west of the Andes, with the exception of P. decumanus. Elsewhere in South 
America, parietal ridges, if present, are usually discontinuous, with the posterior 
component dorsal to and overlapping the anterior component. Specimens having 
a single ridge across each parietal are rare in Amazonian South America and such 
ridges are weakly developed, often inconspicuous. 

The label on the skin of the type has the following information: Proechimys 
semispinosus Tomes; BM 7.1.1.173; Tomes collection; collector L. Fraser 1; 
Gualaquiza; hindfoot 46, ear 21; P. Z. S. London, 1858, p. 548 and 1860, p. 265. 
The catalog number indicates that this specimen was catalogued on | January 
1907, the same date that other specimens Tomes described were catalogued (e.g., 
Marmosa waterhousei, BN 7.1.1.215; Oryzomys albigularis, BM 7.1.1.105; O. 
caliginosus, BM 7.1.1.128; Thomasomys aureus, BM 7.1.1.104). Not all of Fra- 
ser’s material reported on by Tomes was catalogued on that date because the 
types of Oryzomys phaeopus Thomas, 1894 (BM 59.11.1.9), and O. dryas Thom- 
as, 1898 (BM 59.11.1.11), from the same collection had been catalogued in 1859. 
This information suggests that the types of all of the species described by Tomes 
from Fraser’s Ecuadoran collections were among those specimens found at the 
Zoological Society of London. 

The label information that the skin is Fraser’s number | is difficult to interpret. 
The specimen is neither the first that Fraser collected in Ecuador nor the first 
mammal because he said he had ‘“‘three or four Mammalia’ (Fraser 1858a:5942) 
from Cuenca where he began collecting and where Proechimys does not occur. 
One could argue that Fraser’s number | is correct, that he used a separate series 
for each year, and that the holotype of P. se:mispinosus was the first prepared in 
1858 and, therefore, definitely came from Gualaquiza. However, judging by Fra- 
ser’s numbers for birds cited in some of Sclater’s reports (1859c, 1860a, d, e) it 
appears that Fraser used a single series of field numbers during 1858 and 1859 
and presumably from the time he began collecting in 1857. Probably all of his 
specimens or at least all of the vertebrates were included in the single series of 


VOLUME 96, NUMBER | . 141 


numbers. Fraser's number 1705 (the holotype of Oryzomys albigularis) is close 
to Fraser's numbers that Sclater (1860a) cited for birds, also from Pallatanga. 
Therefore, the indication that the type of P. semispinosus was Fraser’s first 
Ecuadoran mammal specimen must be incorrect. 

The only Proechimys whose features resemble those of the type of P. semi- 
spinosus and may be expected to occur in the vicinity of Gualaquiza is P. quad- 
ruplicatus Hershkovitz, 1948. Although some features of P. quadruplicatus are 
similar to those of the type as well as to specimens identified as P. semispinosus 
by Patton and Gardner (1972) from Central America and the Pacific lowlands of 
Colombia and Ecuador, the skulls are clearly different. Contrasted with P. quad- 
ruplicatus, the skull of the type of P. semispinosus has a broader braincase, well- 
developed continuous ridges across the parietals, more posterior termination of 
the nasals in relation to the fronto-maxillary suture, and a less complex pattern 

3-3-3-4 4-4-4-4 
of enamel folds on the cheek teeth 7333 versus 733-5 

So where could Fraser have collected the type of P. semispinosus? Proechimys 
occurs in only four of the localities where Fraser collected: Gualaquiza, Baba- 
hoya, Guayaquil, and Esmeraldas. One of the first places where Fraser worked 
was Gualaquiza where he took the specimen Tomes (1859) reported as “‘Echimys 
cayenensis..’ Tomes’ (1860b) third account listed taxa from several places in 
Ecuador, including Gualaquiza. However, the only Gualaquizan mammals of this 
account were larger species; all of the small mammals Fraser collected at Guala- 
quiza had been covered in Tomes’ first report (1859). There is no evidence that 
Fraser collected mammals in Babahoya or Guayaquil. Esmeraldas was the last 
Ecuadoran locality visited by Fraser and, in my opinion, is the most likely origin 
of the three specimens on which Tomes based his description of P. semispinosus. 
Furthermore, a species of Proechimys characterized by all of the features dem- 
onstrated by the type of P. se:nispinosus is common in the vicinity of Esmeraldas. 
Coincidentally, Esmeraldas was designated by Allen (1913:537) as the type lo- 
cality of Oryzomys caliginosus, the other species Tomes (1860b) described with- 
out locality in his third report. 


in P. quadruplicatus). 


Conclusions 


A review of Fraser’s travels in Ecuador, an examination of the content of 
Tomes’ (1859, 1860a, b) main reports on the mammals Fraser collected, and the 
direct comparison of the type specimen with samples of several species of Proe- 
chimys from Central and South America indicate that Gualaquiza is not the source 
of the type of P. semispinosus. The type locality is here corrected to Esmeraldas, 
Prov. Esmeraldas, on the Pacific coast of Ecuador. 

Much of the confusion regarding the correct allocation of the name P. semispi- 
nosus resulted from the following factors: 1) The presumed loss of the type and 
Thomas’ faith that material from ‘“‘near Bogota’’ was representative of true semi- 
spinosus; 2) the belief (subsequent to the rediscovery of the type) that the type 
locality was Gualaquiza and, therefore, the zoogeographically-based conclusion 
that the name was applicable to populations east of the Andes; and 3) the obvious 
similarity between the type specimen and characteristics seen in populations of 
Proechimys found in Central America and the Pacific lowlands of Colombia and 
Ecuador. 


142 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Because of the correction of the type locality, P. semispinosus calidior be- 
comes a subjective junior synonym of P. semispinosus. The epithets burrus, 
centralis, chiriquinus, colombianus, goldmani, gorgonae, ignotus, panamensis, 
rosa, and rubellus are available as names for populations of P. semispinosus. Of 
these, centralis applies to the populations in Nicaragua, Honduras, and northern 
Costa Rica. The southernmost population of Ecuador should be known as P. 
semispinosus rosa. Allocation of most of the remaining names awaits a taxonomic 
revision of the species. 

Hershkovitz (1948) allied P. ignotus with his P. quadruplicatus group; how- 
ever, several populations of P. semispinosus are superficially similar in dental 
structure to P. quadruplicatus and ignotus is best treated as a subspecies of P. 
semispinosus. 

The Central American distribution of P. semispinosus extends from south- 
eastern Honduras and eastern Nicaragua through Costa Rica and Panama includ- 
ing the Islas las Pearlas (see Hall 1981:873). In South America, the range is west 
of the Andes from the Choco of Colombia (including Isla Gorgona) to south- 
western Ecuador. A record for extreme northwestern Peru (Tumbez, Rio Tum- 
bez; see Thomas 1882:101) has not been confirmed. Those specimens may rep- 
resent P. decumanus Thomas, 1899, which occurs in that region and is sympatric 
with P. semispinosus rosa in southwestern Ecuador (provinces of Guayas and El 
Oro). 


Acknowledgments 


I am grateful to personnel at the British Museum (Natural History), especially 
to Mr. J. E. Hill, for the opportunity to examine type specimens and other ma- 
terial in their care. D. E. Wilson and C. O. Handley, Jr., have provided valuable 
suggestions for improving the manuscript. 


Literature Cited 


Allen, J. A. 1899. The generic names Echimys and Loncheres.—Bulletin of the American Museum 
of Natural History 12:257—264. 

—. 1913. Revision of the Melanomys group of American Muridae.—Bulletin of the American 
Museum of Natural History 32:533—555. 

—. 1916. List of mammals collected in Colombia by the American Museum of Natural History 
Expeditions, 1910—1915.—Bulletin of the American Museum of Natural History 35:191—238. 

Bangs, O. 1901. The mammals collected in San Miguel Islands, Panama, by W. W. Brown, Jr.— 
American Naturalist 35:631-644. 

———. 1905. The Vertebrata of Gorgona Island, Colombia, II]. Mammalia.—Bulletin of the Mu- 
seum of Comparative Zoology 46:89-91. 

Bole, B. P., Jr. 1937. Annotated list of mammals of the Mariato River District of the Azuero 
Peninsula.—Scientific Publications of the Cleveland Museum of Natural History 7:140-188. 

Cabrera, A. 1961. Catalago de los mamiferos de America del Sur.—Revista del Museo Argentino 
de Ciencias Naturales ‘‘Bernardino Rivadavia,’’ Ciencias Zoologicas 4(2):xxii + 309-732, fron- 
tispiece. 

Ellerman, J. R. 1940. The families and genera of living rodents.—British Museum (Natural History), 
London 1|:xxvi + 689 pp. 

Fraser, L. 1858a. [Extracts from letters].—Zoologist 16:5939-5942. 

1858b. Mr. Louis Fraser’s expedition to Ecuador and Peru [extracts from letters].—Zool- 
ogist 16:6158—6160. 
——. 1859a. [Extracts from letters].—Ibis 1:113-114. 


VOLUME 96, NUMBER 1 143 


——. 1859b. [Extracts from a letter].—Ibis 1:208—209. 

——. 1859c. [Extracts from a letter].—Ibis 1:332-333. 

——. 1859d. [Extracts from letters].—Ibis 1:462—464. 

1860. [Extracts from letters].—Ibis 2:192-193. 

Goldman, E. A. 1920. Mammals of Panama.—Smithsonian Miscellaneous Collections 69: 1-309, 1 
map. 

Goodwin, G. G. 1946. Mammals of Costa Rica.—Bulletin of the American Museum of Natural 
History 87:271—474. 

Hall, E.R. 1981. The mammals of North America, Second Ed.—T. Wiley and Sons, New York 2: 

601-1181 + 90 pp. 

, and K. R. Kelson. 1959. The mammals of North America.—Ronald Press, New York 2: 

villi + 547-1083 + 79. 

Handley, C. O., Jr. 1966. Checklist of the mammals of Panama. Pp. 753-795 in R. L. Wenzel and 
V. J. Tipton, eds., Ectoparasites of Panama.—Field Museum of Natural History, Chicago xii + 
861 pp. 

——. 1976. Mammals of the Smithsonian Venezuelan Project.—Brigham Young University Sci- 
ence Bulletin, Biological Series 20: 1-89. 

Hershkovitz, P. 1948. Mammals of northern Colombia. Preliminary report no. 2: Spiny rats (Echi- 
myidae), with supplemental notes on related forms. proceedings of the United States National 
Museum 97:125—140. 

Hollister, N. 1914. Four new Neotropical rodents.—Proceedings of the Biological Society of Wash- 
ington 27:57-60. 

Kellogg, R. 1946. Three new mammals from the Pearl Islands, Panama.—Proceedings of the Bio- 
logical Society of Washington 59:57-62. 

Moojen, J. 1948. Speciation in the Brazilian spiny rats (genus Proechimys, family Echimyidae).— 
University of Kansas Publications, Museum of Natural History 1:301—406. 

Osgood, W. H. 1944. Nine new South American rodents.—Field Museum of Natural History, 
Zoological Series 29:191—204. 

Patton, J. A., and A. L. Gardner. 1972. Notes on the systematics of Proechimys (Rodentia: Echi- 
myidae), with emphasis on Peruvian forms.—Occasional Papers of the Museum of Zoology, 
Louisiana State University 44:1-30. 

Paynter, R. A., Jr., and M. A. Traylor, Jr. 1977. Ornithological gazetteer of Ecuador.—Harvard 
University, Cambridge, Massachusetts. viii + 151 pp. 

Reig, O. A., and M. Useche. 1976. Diversidad cariotipica y sistematica en poblaciones Venezolanas 
de Proechimys (Rodentia, Echimyidae), con datos adicionales sobre poblaciones de Peru y 
Colombia.—Acta Cientifica Venezolana 27:132-140. 

Sclater, P.L. 1859a. List of birds collected by Mr. Louis Fraser at Cuenca, Gualaquiza, and Zamora 

in the Republic of Ecuador.—Proceedings of the Zoological Society of London 1858:449_461. 

1859b. On the birds collected by Mr. Fraser in the vicinity of Riobamba, in the Republic 

of Ecuador.—Proceedings of the Zoological Society of London 1858:549-556. 

——. 1859c. List of the first collection of birds made by Mr. Louis Fraser at Pallatanga, Ecuador; 
with notes and descriptions of new species.—Proceedings of the Zoological Society of London 
1859: 135-147. 

——. 1860a. List of additional species of birds collected by Mr. Louis Fraser at Pallatanga, 
Ecuador; with notes and descriptions of new species.—Proceedings of the Zoological Society 
of London 1860:63-73. 

——. 1860b. List of birds collected by Mr. Fraser in the vicinity of Quito and during excursions 
to Pichincha and Chimborazo; with notes and descriptions of new species.—Proceedings of 
the Zoological Society of London 1860:73-83. 

——. 1860c. List of birds collected by Mr. Fraser in Ecuador, at Nanegal, Calacali, Perucho, 
and Puellaro; with notes and descriptions of new species.—Proceedings of the Zoological 
Society of London 1860:83-97. 

——.. 1860d. List of birds collected by Mr. Fraser at Babahoyo in Ecuador, with descriptions of 
new species.—Proceedings of the Zoological Society of London 1860:272-290. 

——. 1860e. List of birds collected by Mr. Fraser at Esmeraldas, Ecuador, with descriptions of 
new species.—Proceedings of the Zoological Society of London 1860:291-298. 

Tate, G. H. H. 1935. The taxonomy of the genera of Neotropical hystricoid rodents.—Bulletin of 
the American Museum of Natural History 68:295-448. 


144 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Thomas, O. 1882. On a collection of rodents from north Peru.—Proceedings of the Zoological 
Society of London 1882:98-111. 

—. 1896. On new small mammals from the Neotropical region.—Annals and Magazine of 
Natural History (6)18:301-314. 

———. 1897. Descriptions of new bats and rodents from America.—Annals and Magazine of Nat- 
ural History (6)20:544—-553. 

—. 1898. Description of a new Echimys from the neighbourhood of Bogota.—Annals and 
Magazine of Natural History (7)1:243-245. 

———. 1899. Descriptions of new Neotropical mammals.—Annals and Magazine of Natural History 
(7)4:278—288. 

—. 1900. Descriptions of new Neotropical mammals.—Annals and Magazine of Natural History 
(7)5:217-222. 

—. 1911. New rodents from S. America.—Annals and Magazine of Natural History (8)8:250- 
256. 

—. 1914. New Nasua, Lutra, and Proechimys from South America.—Annals and Magazine of 
Natural History (8)14:57-61. 

Tomes, R. F. 1859. Notes on a collection of Mammalia made by Mr. Fraser at Gualaquiza.— 
Proceedings of the Zoological Society of London 1858:546—-549. 

—. 1860a. Notes on a second collection of Mammalia made by Mr. Fraser in the Republic of 
Ecuador.—Proceedings of the Zoological Society of London 1860:211—221. 

—. 1860b. Notes on a third collection of Mammalia made by Mr. Fraser in the Republic of 
Ecuador.—Proceedings of the Zoological Society of London 1860:260-268. 

True, F. W. 1889. On the occurrence of Echinomys semispinosus, Tomes, in Nicaragua.—Pro- 
ceedings of the United States National Museum 1|1:467—468. 


U.S. Fish and Wildlife Service, National Museum of Natural History, Wash- 
ington, D.C. 20560. 


PROC. BIOL. SOC. WASH. 
96(1), 1983, pp. 145-148 


THE ASSIGNMENT OF THE TEXAS TROGLOBITIC WATER 
SLATER CAECIDOTEA PILUS TO THE GENUS 
LIRCEOLUS, WITH AN EMENDED 
DIAGNOSIS OF THE GENUS 
(CRUSTACEA: ISOPODA: ASELLIDAE) 


Julian J. Lewis 


Abstract.—Caecidotea pilus is redescribed and assigned to Lirceolus. This ge- 
nus, now containing two species, remains endemic to Texas. The addition of C. 
pilus requires emendation of Lirceolus to broaden the concept of the genus, 
especially in the characteristics of the mouthparts. 


Until recently, all of the known subterranean asellid isopods of Texas were 
assigned to the widespread genus Caecidotea. In 1976 Bowman and Longley 
redescribed Caecidotea smithii from newly collected specimens from the artesian 
well at San Marcos, Texas. This species possessed a number of unusual mor- 
phological characters that persuaded Bowman and Longley to place it in a new 
genus, Lirceolus, so named for the similarity of the third pleopods of Lirceus 
and Lirceolus. 

Lewis (1982) pointed out the morphological similarities of some of the other 
Texas asellids to Lirceolus and suggested the possibility that under scrutiny, 
additions might be made to the genus. Of the three other troglobitic asellids from 
Texas (all described by Steeves 1968), Caecidotea reddelli is clearly assigned to 
the correct genus. On the other hand the illustrations of C. pilus and C. bisetus 
by Steeves (1968) were suggestive of Lirceolus in the unarmed gnathopod, elon- 
gate first pleopod, and the sparsely setose exopod of the second pleopod. When 
examined, C. bisetus was found to have the usual Caecidotea-type transverse 
suture across the exopod of the third pleopod. Caecidotea pilus possessed not 
only an oblique suture, but other characteristics which necessitate its transfer- 
ence to Lirceolus. However, some of the unusual characteristics of Lirceolus’ 
smithii included in the diagnosis of the genus by Bowman and Longley (1976) do 
not apply to L. pilus; hence an emended diagnosis is provided here. 

Several illustrations are included herein to support the placement of L. pilus 
in Lirceolus. Steeves (1968) illustrated the male gnathopod, pleopod | and pleo- 
pod 2, plus the endopod tip of the latter. Except where new details have been 
revealed, Steeve’s drawings adequately characterize the species and are not re- 
peated here. 


Lirceolus Bowman and Longley, 1976 


Diagnosis.—Eyeless, unpigmented, maximum length about 4 mm. Head with- 
out lateral incisions or rostrum. Mandible with 3-merous palp. Maxilla 1, outer 
lobe with 10-13 spines; inner lobe with 5-8 plumose setae. Pereopod | propodus 
palm without processes. Pleopod | slender, elongate, distal segment oval with 
sparse non-plumose setation. Pleopod 2, exopod proximal segment produced me- 


146 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


dially over distal segment; distal segment without catch lobe, with longitudinal 
furrow on posterior surface and 0-1 setae on distal margin. Endopod with short 
basal spur, basal apophysis about as long as basal spur, labial spur absent. Pleo- 
pod 3 exopod with oblique suture. Pleopods 4 and 5 with exopod and endopod 
partly or totally fused. 

Type-species.—Asellus smithii Ulrich, 1902. 


Lirceolus pilus (Steeves) 
Fig. | 


Asellus pilus Steeves, 1968:188.—Reddell and Mitchell, 1969:8, 43.—Reddell, 
1970:396.—Fleming, 1973:295 (in list), 297 (in key). 

Conasellus pilus (Steeves).—Henry and Magniez, 1970:356.—Mitchell and Red- 
diel, ISHiss >. 


Material examined.—YEXAS: Medina Co., Valdina Farms Sinkhole, 15 mi. 
N. Sabinal, 12 Jan 1963, leg. J. Reddell, D. McKenzie, J. Porter, holotype d 
(USNM 119593), allotype 2 (USNM 119594), 15, 12 paratypes (USNM 119595). 

Description.—Maximum length 3.0 mm (allotype), body slender, about 4x as 
long as wide. Coxae visible in dorsal view. Head about 2x as wide as long. 
Pleotelson about 1.3x as long as wide, sides subparallel, caudomedial lobe not 
evident. 

Antenna | flagellum of about 5 segments, esthetes on distal 3 segments (d 
paratype) or 1-0-1 (2 paratype). Mandibles with 4-cuspate incisors and lacinia 
mobilis, palp with few plumose setae on segments 2 and 3. Maxilla 1 inner lobe 
with 5 plumose setae; outer lobe with 13 robust spines. Maxilliped with 3 reti- 
nacula. 

Pereopod | propod about 3.0x as long as wide in 6, 2.4x in 9. Pereopod 4 
sexual dimorphism slight, carpus of holotype 3.3 x as long as wide, allotype 3.5x; 
dactyl with accessory unguis. 

Pleopod | with 3 retinacula; exopod about 1.4x< length of protopod. Pleopod 
2, exopod distal segment with | seta; endopod tip with endopodial groove ter- 
minating in decurved beak-shaped process. Pleopod 3 as figured. Pleopods 4 and 
5 exopods with single oblique suture. 

Distribution.—Known only from the type-locality, Valdina Farms Sinkhole. 
The assignment of this species to Lirceolus extends the range of the genus a short 
distance to the west, but it remains endemic to the Balcones Fault Zone of Texas. 

Relationships.—Lirceolus pilus is obviously closely related to L. smithii both 
morphologically and geographically. The greatest differences between the 2 species 
lies in the structure of their mouthparts: 


L. smithii L. pilus 
mandibles: 
lacinia/incisors 2-3 cuspate 4-4 cuspate 
maxilla 1 
outer lobe 10 spines 13 spines 
inner lobe 8 setae 5 setae 


The generic relationships of Lirceolus remain obscure, although the addition 


VOLUME 96, NUMBER 1 147 


Fig. 1. Lirceolus pilus: A from allotype, B, C, F, G from ¢ paratype, others from 92 paratype. 
A, Pereopod 4 dactyl; B, Pleopod 2, exopod; C, Same, endopod tip; D, Pleopod 5; E, Incisor and 
lacinia, left mandible; F, incisor, right mandible; G, Mandibular palp; H, Antenna 1; I, Maxilla 1, 
inner lobe; J, Same, outer lobe; K, Pleopod 4; L, Pleopod 5. 


of L. pilus lends some insight. The mouthparts of L. pilus are very similar to 
those of troglobitic Caecidotea both morphologically and meristically. However, 
the oblique suture of pleopod 3 clearly separates Lirceolus from Caecidotea. 
Lirceolus can be separated from Lirceus by the lack of a rostrum and the lateral 
incisions of the head, present in most species of Lirceus. A fusion of the endopod 
and exopod of pleopods 4 and 5 as in L. smithii is unknown in both Caecidotea 
and Lirceus, although in Calasellus, pleopod 5 exopod is greatly reduced or 
absent (Bowman 1981). 

The fusion of the pleopods reported in L. smithii by Bowman and Longley 
(1976) was difficult to ascertain in L. pilus. In the female paratype the pleopods 


148 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


appeared fused similar to those reported for L. smithii, but the fusion looked 
much less extensive in the male paratype. The unusual fleshy consistency of the 
pleopods in Lirceolus pilus (and Lirceolus in general) precludes an adequate 
description from the few specimens available. Moreover, all specimens were 
incomplete, lacking the flagella of the second antennae, the uropods, and most 
of the pereopods. Of the two males and two females in the collection, only one 
individual of each sex retained a single fourth pereopod to compare for sexual 
dimorphism. 


Acknowledgments 


I thank Dr. Thomas E. Bowman for the loan of specimens from the collections 
of the Smithsonian Institution, and for reading this manuscript. This paper is 
Contribution #206 (New Series) of the Department of Biology, University of 
Louisville. 


Literature Cited 


Bowman, Thomas E. 1981. Calasellus longus, a new genus and species of troglobitic asellid from 
Shaver Lake, California (Crustacea: Isopoda: Asellidae).—Proceedings of the Biological So- 
ciety of Washington 94(3):866—872. 

, and Glenn Longley. 1976. Redescription and assignment to the new genus Lirceolus of the 

Texas troglobitic water slater, Asellus smithii (Ulrich) (Crustacea: Isopoda: Asellidae).—Pro- 

ceedings of the Biological Society of Washington 88(45):489-496. 

Fleming, Laurence E. 1973. The evolution of the eastern North American isopods of the genus 
Asellus (Crustacea: Asellidae). Part I1.—International Journal of Speleology 5:283-—310. 
Henry, Jean-Paul, and Guy Magniez. 1970. Contribution a la systématique des Asellides (Crustacea 

Isopoda).—Annales de Spéléologie 25(2):335—367. 

Lewis, Julian J. 1982. A diagnosis of the Hobbsi Group, with descriptions of Caecidotea teresae, 
n. sp. and C. macropropoda Chase and Blair (Crustacea: Isopoda: Asellidae).—Proceedings 
of the Biological Society of Washington 95(2):338-346. 

Mitchell, Robert W., and James R. Reddell. 1971. The invertebrate fauna of Texas caves. Pp. 35— 
91 in Ernest L. Lundelius and Bob H. Slaughter, (eds.), Natural History of Texas Caves. Gulf 
Natural History, Dallas. 

Reddell, James R. 1970. A checklist of the cave fauna of Texas. IV. Additional records of Inver- 

tebrata (exclusive of Insecta).—Texas Journal of Science 21(4):389-415. 

, and Robert W. Mitchell. 1969. A checklist and annotated bibliography of the subterranean 

aquatic fauna of Texas.—Texas Technological College, Water Resources Center, Special Re- 

port 24:1—48. 

Steeves, Harrison R. III. 1968. Three new species of troglobitic asellids from Texas.—American 
Midland Naturalist 79(1): 183-188. 

Ulrich, Carl J. 1902. A contribution to the subterranean fauna of Texas.—Transactions of the 
American Microscopical Society 23:83—101. 


Department of Biology and Water Resources Laboratory, University of Louis- 
ville, Louisville, Kentucky 40292. 


PROC. BIOL. SOC. WASH. 
96(1), 1983, pp. 149-153 


CAECIDOTEA FONTICULUS, THE FIRST TROGLOBITIC 
ASELLID FROM THE OUACHITA MOUNTAINS 
(CRUSTACEA: ISOPODA: ASELLIDAE) 


Julian J. Lewis 


Abstract.—Caecidotea fonticulus, a member of the Hobbsi Group, is the first 
troglobitic isopod to be described from the Ouachita Mountains. This species 
appears to be most closely related to two species from the Ozarks, C. spatulata 
and C. fustis. 


The species described herein is the first troglobitic asellid to be found from the 
Ouachita Mountains of central Arkansas and adjacent Oklahoma. Another species 
occurring in the Ouachitas, Caecidotea oculata, has reduced eyes and pigmen- 
tation, but is reported only from epigean habitats (Mackin and Hubricht 1940). 
The discovery of this zoogeographically interesting new asellid was made possible 
by an extensive ongoing survey of Ouachita springs and their fauna by Dr. Henry 
W. Robison of Southern Arkansas University. 


Caecidotea fonticulus, new species 
Figs. 1-2 


Material examined.—ARKANSAS: Polk Co., Abernathy Spring, 0.8 mi. W of 
Polk-Montgomery county line, on north side of Highway 8 (shown on U.S.G.S. 
Big Fork, Arkansas 7.5 minute quadrangle), 16 June 1979, H. W. Robison, 136 
36 2 (USNM 191132); same locality, H. W. Robison, 21 May 1980, 16 36, 14 
2 2 (USNM 191129); same locality, H. W. Robison, 16 Apr 1982, 29 62 (USNM 
191133); same locality, Julian J. Lewis and Teresa M. Lewis, 31 May 1981, 9 
366,10 22 (USNM 191131). 

A 5.0 mm ¢ from the 31 May 1981 collection is the holotype (USNM 191128), 
the other specimens are paratypes. All of the material examined has been de- 
posited in the National Museum of Natural History, Smithsonian Institution. 

Description.—Eyeless, unpigmented. Longest ¢ 6.0 mm, longest 2 5.0 mm; 
body slender, about 6.1 as long as wide. Head about 1.5x< as wide as long, 
anterior margin concave, postmandibular lobes moderately produced. Pleotelson 
about |.2x as long as wide, sides subparallel, caudomedial lobe produced. 

Antenna | reaching midlength of last segment of peduncle of antenna 2, fla- 
gellum with up to 8 segments, esthete formula to 6-0, occasional segments with 
2 esthetes. Mandibles with 4-cuspate incisors and lacinia mobilis; palp with plu- 
mose setae in rows on distal segments. Maxilla 1, outer lobe with 13 robust spines, 
inner lobe with 5 plumose setae. Maxilliped with 5-6 retinacula. 

Male pereopod | propus about 1.4 as long as wide; palmar margin with 2 
short spines proximally, high subtriangular median process separated by U-shaped 
cleft from similar lower distal process; dactyl flexor margin without process, 
undulating, with small spines. Female pereopod | about 2x as long as wide, 


150 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 1. Caecidotea fonticulus: a—h from ¢ paratypes, i from @ paratype: a, Habitus, dorsal; b, 
Incisor and lacinia, left mandible; c, Pereopod 1; d, Antenna 1, distal segments; e, Mandibular palp; 
f, Maxilla 1, inner lobe; g, Same, outer lobe; h, Pereopod 4, distal segments; i, Same. 


VOLUME 96, NUMBER 1 151 


Fig. 2. Caecidotea fonticulus: a-i from 3 paratypes, j from @ paratype: a, Pleopod 1; b, Pleopod 
2; c-e, Same, endopod tip, anterior, posterior, lateral views; f, Pleopod 3; g, Pleopod 4; h, Pleopod 
5; 1, Uropod; j, Same. 


152 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


propus without processes. Pereopod 4 of ¢ and @ similar, carpus about 2.4x as 
long as wide. 

Male pleopod | longer than pleopod 2; protopod about 0.6 length of exopod, 
with 4—5 retinacula; exopod about 2.3x as long as wide, rounded distal margin 
with 1-2 elongate plumose setae, concave lateral margin without plumose setae. 
Male pleopod 2 exopod, proximal segment with 4 lateral setae, distal segment 
with about 8 long plumose setae along distal margin; endopod with distinct basal 
apophysis, tip with 4 processes: (1) caudal process broadly rounded, with scal- 
loped surface, (2) lateral process recurved, extending slightly beyond caudal pro- 
cess, (3) cannula conical, truncate, directed distally, endopodial groove mostly 
obscured except in lateral view, and (4) mesial process low, forming a shelf across 
base of cannula in anterior view. Pleopod 3 exopod distal margin with sparse 
non-plumose setae. Pleopod 4 exopod with single sigmoid suture trifurcating to 
notches in lateral margin, seta present or absent, many setules present. Pleopod 
5 with 2 transverse sutures. Uropods about 1.4 length of pleotelson in d, 0.7 
im ©. 

Etymology.—The name, proposed as a noun, is derived from the Latin “‘fon- 
ticulus’’ (a little fountain or spring) in reference to the type-locality of the species. 

Distribution.—Known only from the type-locality, Abernathy Spring. The spring 
flows from a tile pipe sunken lengthwise into the ground, giving the fountain-like 
appearance to which the specific name refers. After a few meters the spring 
stream joins another stream, Big Fork. Two species of asellids are found under 
rocks and in aquatic mosses in the spring stream, C. fonticulus and Lirceus 
ouachitaensis. This is the first record of this Lirceus from Arkansas, although 
Mackin and Hubricht (1938) reported it from the adjacent part of the Ouachitas 
in Oklahoma. Specimens of this asellid have also been deposited in the Smith- 
sonian Institution (USNM 191130). The water from Abernathy Spring averages 
about 16°C, pH 7.0, conductivity 144 umhos and alkalinity 70 mg/ml CaCO, 
(Robison, in litt.). 

Relationships.—Caecidotea fonticulus can be readily assigned to the Hobbsi 
Group (as defined by Lewis 1982) by the following combination of male charac- 
ters: pleopod | longer than pleopod 2, distal margin with elongate plumose setae, 
and pleopod 2 endopod tip with bluntly conical, distally directed cannula. Spe- 
cifically, the morphology of the endopod of C. fonticulus resembles that of two 
Ozark species, C. spatulata and C. fustis (Mackin and Hubricht 1940; Lewis and 
Bowman 1981; Lewis 1981). All three species have in common a high, somewhat 
digitiform lateral process that is slightly recurved, a broad mesial process that 
obscures the base of the cannula, and a broad caudal process. The palmar margin 
of the gnathopod propus bears a triangular median process in these species, but 
bicuspid distal processes in C. spatulata and C. fustis, instead of the triangular 
distal process of C. fonticulus. The Ozark species are readily separated from C. 
fonticulus by their numerous elongate plumose setae along the distal margin of 
the third pleopod. In some populations of C. spatulata and C. fustis vestigial 
eyes and pigmentation are present, a characteristic not exhibited by any of the 
specimens of C. fonticulus examined. 


VOLUME 96, NUMBER 1 153 


Acknowledgments 


I thank Dr. Henry W. Robison for visiting Abernathy Spring on several oc- 
casions to collect specimens, and for supplying me with detailed information on 
the spring. Dr. Thomas E. Bowman called my attention to the first specimens of 
this new species (which had been deposited in the collection of the Smithsonian 
Institution by Dr. Robison) and read yet another manuscript. My wife, Teresa 
M. Lewis, provided field assistance on a 1981 collecting trip through the Ouachi- 
tas and Ozarks, funded by.a grant from the Graduate School of the University 
of Louisville. This paper is contribution #208 (New Series) from the Department 
of Biology, University of Louisville. 


Literature Cited 


Lewis, Julian J. 1981. Caecidotea salemensis and C. fustis, new subterranean asellids from the 
Salem Plateau (Crustacea: Isopoda: Asellidae).—Proceedings of the Biological Society of 
Washington 94(2):338—346. 

——. 1982. A diagnosis of the Hobbsi Group, with descriptions of Caecidotea teresae, n. sp., 

and C. macropropoda Chase and Blair (Crustacea: Isopoda: Asellidae).—Proceedings of the 

Biological Society of Washington 95(2):81—94. 

, and Thomas E. Bowman. 1981. The subterranean asellids (Caecidotea) of Illinois (Crus- 

tacea: Isopoda: Asellidae).—Smithsonian Contributions to Zoology 335, 66 pp. 

Mackin, J. G., and Leslie Hubricht. 1938. Records of distribution of species of isopods in central 

and southern United States, with descriptions of four new species of Mancasellus and Asellus 

(Asellota, Asellidae).—American Midland Naturalist 19:628—637. 

, and 1940. Descriptions of seven new species of Caecidotea (Isopoda, Asellidae) 

from the central United States.—Transactions of the American Microscopical Society 59(3): 

383-397. 


Department of Biology and Water Resources Laboratory, University of Louis- 
ville, Louisville, Kentucky 40292. 


PROC. BIOL. SOC. WASH. 
96(1), 1983, pp. 154-159 


PSYCHRONAETES HANSENI, A NEW GENUS AND 
SPECIES OF ELASIPODAN SEA CUCUMBER 
FROM THE EASTERN CENTRAL PACIFIC 
(ECHINODERMATA: HOLOTHUROIDEA) 


David L. Pawson 


Abstract.—A new genus and species in the elasipodan family Laetmogonidae 
is described. Distinguishing features include large size (largest specimens prob- 
ably exceed 30 cm in length), presence of a pronounced anterior “‘head’’ region 
with large, irregularly sized papillae, presence in the bodywall of wheels of only 
one type, and absence of circumoral papillae. The genus is known so far only 
from the eastern central Pacific. A revised key to genera in the family Laetmo- 
gonidae is presented. 


During a benthic sampling and photographic survey of a large area of the east- 
ern central Pacific, the Ocean Mineral Company collected specimens of a rea- 
sonably common, large, elasipodan holothurian, which proved to represent a new 
genus. Seafloor photographs of the animal in situ were also obtained. Specimens 
and photographs were entrusted to me for study. 

Hansen (1975) revised the elasipodan family Laetmogonidae, and it now com- 
prises four genera. The new genus described here is distinctive in several impor- 
tant respects, and appears to bear no close relationship to any other genera in 
the family. 


Order Elasipodida Theel, 1882 
Family Laetmogonidae Ekman, 1926 
Psychronaetes, new genus 


Diagnosis.—Body fusiform, maximum length approximately 30 cm. Midventral 
radius naked, lateral ventral radii each with approximately 15 triangular tubefeet. 
Dorsal radii each with approximately 30 papillae. Dorsal papillae forming an 
irregular fringe around anterior dorsal end of body. Tentacles 15, no circumoral 
papillae. Ossicles in bodywall wheels of one type, usually 50-60 um in diameter, 
usually with 9-12 spokes. 

Type-species.—Psychronaetes hanseni, new species 

Etymology.—The genus-name is of masculine gender, derived from Greek, 
psychros cold, and naetes inhabitant. The species is named in honor of Dr. Bent 
Hansen of the Universitetets Zoologiske Museum, Copenhagen, in acknowledg- 
ment of the valuable contributions he has made to our knowledge of the deep- 
sea holothurian fauna. 

Remarks.—This new genus can be distinguished from others in the family Laet- 
mogonidae as follows. The key is a modified version of that given by Hansen 
(1975). 


VOLUME 96, NUMBER 1 155 


Key to genera of family Laetmogonidae 


PECireiiMmnocal paprllacuprKeSseMmt a... +> see 4.6 ok Benthogone Koehler, 1896 
CGrHemmonralpapillac abSeMiinc. 255-8 Wee ses ects ila WEN God oot wonad De kore 2 
2. Midventral tubefeet present. Body wall ossicles wheels with marginal 
Ke CHIME SANE t te acne tet wy alaalia eltodaalinne on ateratels Pannychia Theel, 1882 
Midventral tubefeet absent. Wheels lack marginal teeth ................ 3 


3. Ventrolateral papillae present, fused to form continuous brim around body 
TAS corse eas, wie ter Nee Scene. eae Apodogaster Walsh, 1891 
— Ventrolateral papillae absent. Brim, when present, composed of fused 


WemMURG a TenAImCUD Che iu ryaty «main etek arama tens nel UAC. SU Lee ae aaty Westar 3 4 
4. Body fusiform, up to 30 cm long, with pronounced anterior “‘head‘‘ region 
with irregular fringe of dorsal papillae ........ Psychronaetes, new genus 


— Body more or less cylindrical, usually considerably less than 20 cm long, 
lacking pronounced anterior “‘head’’ region. 
No anterior fringe of dorsal papillae ............ Laetmogone Theel, 1879 


Psychronaetes hanseni, new species 
ES Se lee 


Diagnosis.—As for genus. 

Material examined.—HOLOTYPE, USNM E27566, total length approximately 
240 mm, greatest width approximately 60 mm; PARATYPE USNM E27567, total 
length approximately 70 mm, greatest width approximately 25 mm. Type-locality, 
eastern central Pacific, Clarion-Clipperton Fracture Zone, West of 120°W, depth 
4800-5200 meters, collected by Ocean Minerals Company. 

Description.—Color in alcohol variegated dark purple, lighter in smaller spec- 
imens. Body fusiform, tapering anteriorly and posteriorly (Fig. 1). Pronounced 
‘neck’ anteriorly, approximately 15% of body length from anterior end. Head 
region with mouth, ventrally directed, and tentacles. No circumoral papillae. 
Tentacles 15, with short stems and large elongate oval discs 9 x 5 mm in holo- 
type, with long axis of disc directed towards mouth. Anus posterodorsal. Body- 
wall firm, leathery, not gelatinous. 

Midventral radius naked. Lateral ventral radii each with 15 large and conspic- 
uous triangular tubefeet, each approximately 10 mm long and 8 mm wide at base. 
Feet evenly scattered along entire lateral ventral radii. Dorsal surface with 2 
poorly defined rows of numerous papillae, approximately 30 in each row, papillae 
apparently arising from medial side of each dorsal radius. On head region, papillae 
become enlarged, forming an irregular fringe around anterior dorsal end of body. 
Fringe contains approximately 15 papillae in larger specimens, largest papillae 
approximately 15 mm long and 9 mm wide at base. 

Ossicles in bodywall exclusively wheels, greatly variable in size (Table 1). 
Wheels having smooth rim, short spokes (Table 2) and large central region; at 
center of wheel, raised hub made up of 4 struts. Dorsal bodywall wheels are 
slightly larger in average diameter and tend to have a greater number of spokes 
than wheels from ventral bodywall. Entire wheel strongly concave (Fig. 2a). 
Wheels present everywhere in bodywall, and also in tubefeet and papillae. Ten- 
tacles contain wheels, also rods (Fig. 2c) of greatly variable size (150 um to 1 


156 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


VOLUME 96, NUMBER 1 157 


Fig. 2. Psychronaetes hanseni, ossicles: A, Wheels from bodywall; B, Stage in development of 


bodywall wheel; C, Rods from tentacles. 


— 
Fig. |. Upper and lower, specimens of Psychronaetes hanseni on the seafloor in the vicinity of 


the type-locality. Dark-colored objects are manganese nodules. Each specimen is approximately 25 
cm long. Note anterior ““head’’ region with papillae, and, on upper specimen, the double row of 


dorsal papillae. 


158 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Table 1.—Diameter of wheel ossicles from bodywall. 
Dorsal wheels (n = 50) : Ventral wheels (n = 50) 
Size ranges (jm) Size range 51-80 um Size range 48-76 um 
46-50 0 0% 3 6% 
51-55 14 28% 10 20% 
56-60 18 36% 23 46% 
61-65 3 6% 4 8% 
66-70 9 18% 4 8% 
71-75 5) 10% 4 8% 
76-80 I 2% D 4% 
Table 2.—Number of spokes in wheel ossicles from bodywall. 
Dorsal (n = 50) Ventral (n = 50) 
Number of spokes Number of wheels Number of wheels 
8 0 0% 8 16% 
9 3 6% 16 32% 
10 10 20% 17 34% 
11 ]2 24% 6 12% 
12 23 46% 3 6% 
13 2 4% 0 0% 


mm long), usually slightly curved, often with slightly prickly ends. Developmental 
stages of wheels common in bodywall and tentacles (Fig. 2b). At an early stage 
of development, before wheel rim has formed, central 4-strut hub is underlain by 
tripartite piece; this piece eventually disappears, and is seldom seen in fully 
developed wheels. 

Habitat and ecological aspects.—The sediments on which Psychronaetes han- 
seni occurs are commonly Quarternary siliceous oozes and siliceous clays con- 
sisting of clay minerals and siliceous microfossils. These sediments are fine, 65% 
of the material having a particle size of less than 4 wm. Preliminary analyses of 
seafloor photographs indicate that in three discrete sampling areas, population 
densities of approximately one individual of Psychronaetes hanseni per 1000 m? 
occur. 

Remarks.—Psychronaetes hanseni is among the largest known elasipodan ho- 
lothurians. It is surprising that this distinctive species was not encountered during 
the ‘Albatross’ expeditions in the eastern tropical Pacific (Clark 1920). Its ap- 
parent absence from “‘Challenger** and other collections is perhaps due to its 
restricted distribution pattern. Certainly, this species can be identified in seafloor 
photographs, and no specimens representing this species have as yet been rec- 
ognized in collection of several thousands of photographs from the southern Pa- 
cific and southern Atlantic Oceans examined by the author. 


Acknowledgments 


I am grateful to Dr. Charles Morgan of the Lockheed Ocean Laboratory for 
giving me access to specimens and photographs of this new holothurian, and for 
helping me in many other ways. Dr. Jean Nichols and Ms. Pamela Springer of 


VOLUME 96, NUMBER 1 159 


the same institution were also very helpful in providing habitat data and selected 
photographs. 


Literature Cited 


Clark, H. L. 1920. Holothurioidea—Memoirs of the Museum of Comparative Zoology 39(4):118— 
154. 

Hansen, B. 1975. Systematics and biology of the deep-sea holothurians. Part 1. Elasipoda.—Gal- 
athea Report 13:1—262. 


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


PROC. BIOL. SOC. WASH. 
96(1), 1983, pp. 160-177 


LIFE DIAGRAM PATTERNS IN BENTHIC 
POLYCHAETES 


Kristian Fauchald 


Abstract.—Life diagrams in polychaetes can be grouped into three patterns. 
One pattern, including annual species, shows large reproductive efforts, small 
eggs and planktotrophic larvae. A second pattern, including the perennial species, 
shows low reproductive efforts, moderately large to large eggs, and non-plank- 
totrophic development. The third pattern includes the multi-annual species, shows 
high reproductive effort, moderately large eggs, and non-planktotrophic devel- 
opment. These patterns may be in part systematically related to the environment 
in which the different species of polychaetes are found, but also appear related 
to a potential need for maintaining small cohesive reproductive units. 


The study of life diagrams can be done from several points of view. A veritable 
jungle of modelling studies of various phases of life diagrams has been published 
(reviewed by Menge 1975, and Stearns 1976). General problems with modelling 
have been reviewed recently by Pielou (1981) and will not be considered here. It 
appears intuitively obvious that a certain factual base of observations must be 
available to model-builders so that models represent something known to occur 
with a frequency higher than zero in nature. Conceptual models do not necessarily 
have to be testable in toto, but at least certain facets should be available for 
testing. The lifespan of an individual polychaete may be as short as a few weeks 
(Ophryotrocha spp.) or last several years (Eunice spp.). A basic assumption of 
this paper is that knowledge of the lifespan of any polychaete species can be used 
to predict features in the life diagram of that species, and conversely, that infor- 
mation about certain life diagram features, such as brood size or egg size, can be 
used to predict general life habits and longevity of a given species. 

The primary object of this study is to review the few cases for which infor- 
mation is available to test the assumption. A secondary purpose is to review 
some of the literature on the topic. Thirdly, various predictions are made about 
unknown features of life diagrams for selected species based on the available 
information, and suggestions are made as to how these predictions can be tested. 

The terms life cycle or life history are usually defined as a description of the 
life of an individual starting at release from the parental organism and ending at 
death. Both terms have unfortunate implications. For example, the sequence of 
events is only in a formal sense cyclic since all individuals that go through the 
‘“‘cycle’’ are different (see Wiley and Brooks 1982:1—3), except perhaps in a few 
cloning forms and even in these, postparturition selection may “‘change’’ the 
clones from one release (spawning) episode to the next. The term cycle implies 
a static, non-evolutionary point of view. This concept can be useful, but will aid 
little in explaining differing powers of ecological and geographical dispersal. The 
term “‘history,’’ as used in such combinations as natural history, implies a de- 
scription of past events with little heuristic power. The term has also frequently 
been used to cover the first part of ontogenesis up to reproductive maturity. The 


VOLUME 96, NUMBER 1 161 


information yielded by ontogenetic study of a given species has turned out to be 
useful as a predictor of ontogenesis in phylogenetically or ecologically related 
taxa. The word diagram lacks some of the problematic implications of the two 
other terms. It is clearly a theoretical, formal term and implies a level of abstrac- 
tion useful for making testable predictive statements. The above terminological 
problems were first pointed out to me by Mr. Bill Kennedy. 

Most studies on polychaete life diagrams have focused on a single species, or 
one part of the diagram has been studied for several, usually taxonomically related 
species (cf. papers cited in Schroder and Hermans 1975). Reviews, such as the 
very extensive one issued by Schroder and Hermans (1975) have usually focused 
on detailing the complex factual information necessary for a more theoretical 
approach. In the review quoted, however, the format did not allow much freedom 
to explore the theoretical implications. 

Trendall (1982) pointed out that in mosquito fish, the different life diagram traits 
were not necessarily correlated from one locale to another. For this reason, life 
diagrams must be characterized based on a single population; possibly, for most 
species, differences between populations in life diagram features will be less than 
differences between species, but this cannot be assumed a priori. The path of 
generalizations about life diagrams leads then from a study of single populations, 
via comparisons of diagrams within species, to that between species and between 
higher taxa. In this study it is assumed that differences among populations are 
less than differences among species. The study is based on information about 
single populations, but it is assumed that the diagram found for a population is 
characteristic of the species as a whole. 

The sequence of events in the life of a number of specimens of a single species 
from fertilization to death may be generalized into a life diagram for that species. 
A comparison of life diagrams for several species may be generalized into a 
presumably limited number of life diagram patterns. A life diagram pattern 1s 
thus a second-level abstraction, rather than a primary abstraction. The publ‘ched 
literature makes it amply clear that a distinction between the two levels of ab- 
Straction has not been made. Life diagram patterns do not necessarily follow 
taxonomic lines, even if a primary testable prediction for any species would be 
that its life diagram ought to be similar to that of a related species. Life diagram 
patterns may follow habitat requirements and a secondary set of hypotheses for 
a given species might be associated with this feature. The usefulness of separating 
between these two levels of abstraction thus lies in the possibilities it gives for 
making different sets of predictions. 

Any event in the life diagram of any species can only be wholly understood in 
connection with all other events in the life of each organism. In other words, the 
life diagram represents a set of co-adapted features consisting of several partially 
distinguishable stages or phases, where any phase may be studied separately, but 
where its significance may be understood only as part of the whole diagram. 


Life Diagram Events 


The life of any organism is a continuum. For the purposes of analysis, this 
continuum may be separated into a series of definable events, with the caveat 
that each of the defined events usually is not distinctly separable from the pre- 
vious event or from the following event but forms part of the continuum. For the 


162 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


purpose of this discussion nine distinct life diagram events are recognized and 
defined. The physical location of each event is of primary. importance, not least 
since the location is frequently associated with one or another axis of the life 
diagram (e.g., small egg-size is frequently correlated with pelagic early devel- 
opment; for example see Thorson 1950). 

Spawning and fertilization.—Spawning may take place in the open water, in a 
burrow or tube, or into an eggmass, or the eggs may be retained in the body 
cavity of the female so that internal development takes place (Smith 1950). Eggs 
spawned inside a tube may be grouped in capsules, as in spionids (Blake 1969), 
or they may occur singly in a string, or stored in some other fashion (Knight- 
Jones et al. 1972). Egg sizes may vary from about 30 um to about 2-3 mm in 
diameter. The eggs are usually somewhat flattened or disc-shaped. Fertilization 
usually appears closely associated with the spawning of the eggs and is assumed 
to take place when the eggs are released. The process has been poorly studied 
and nothing is known of the location of fertilization for most species. 

Pre-release development.—The development prior to release of the larvae is 
usually based on energy contained within the egg-membrane, this part of the life 
diagram being fueled by yolk supplied by the parent. Early development usually 
takes place wherever fertilization took place. Several species do not have true 
larvae; the phase is here taken to include a shorthand description of the early 
development prior to the release from the egg-membrane, or to the transformation 
of that membrane into part of the larval or juvenile cuticle. 

Larval stage.—This stage includes a number of pre-metamorphic, post-hatching 
developmental phases, whether they are true trochophores or not. These phases 
may be feeding larvae, in which case they may feed in any one of several different 
environments. However, most feeding polychaete larvae are planktotrophic in 
that they feed pelagically, usually on phytoplankton. Other larvae, such as the 
encapsulated spionid larvae, are adelphophagic, feeding on eggs present in the 
same capsule. In this particular kind of parental provisioning, a series of eggs are 
laid within a single capsule, but only one or a few larvae survive to hatching 
(Blake 1969). 

Usually, but not invariably, feeding larvae are pelagic while non-feeding larvae 
keep close to or on the bottom (Schroder and Hermans 1975). Non-feeding larvae 
are usually morphologically rather different from the characteristic trochophore, 
but vary widely among themselves. For example, the barrel-shaped larvae in the 
super-family Eunicea have short apical tufts of cilia and a broad band of short 
cilia around the middle. At the time of release from the parental tube, they are 
usually poorly differentiated internally (Akesson 1967). Orbiniid larvae are com- 
pletely covered with short cilia (Anderson 1959, 1961, 1966), and recognition of 
the prototrochal cilia can be difficult. 

The length of the trochophoral stage varies tremendously. Certain spirorbids 
are competent to settle at the time of release (Beckwitt 1979). In contrast, certain 
sabellariids may remain pelagic for several months (reviewed by Mauro 1975). 
Perhaps the most common pattern is for the larvae to become competent to settle 
fairly rapidly, for example after a 10-day stay in the plankton, with the heavy 
settlement following after an additional 10 days in the plankton (Thorson 1946), 
but the larvae may be able to stay in the plankton for as much as several weeks 
(Wilson 1968). 


VOLUME 96, NUMBER 1 163 


Metamorphosis.—Most polychaetes undergo a gradual metamorphosis in which 
the posterior half of the larva elongates and becomes segmented, with setae 
forming in each segment (Schroder and Hermans 1975). Two to three recogniz- 
able setigers are usually formed before juveniles settle out of the plankton, but 
in some polychaetes, especially spionids,; as many as 20 setigers may be present 
while juveniles are still in the plankton (Blake 1969, Hannerz 1956). 

The pattern of metamorphosis is rather similar in both feeding and non-feeding 
larvae, but is usually less drastic in the latter since they usually do not form 
complex larval structures that have to be modified for adult use. In most species 
the start of the metamorphic events appears to be under endogenous control in 
that formation of segments and setae start while individuals are still in the larval 
habitat. The process is however frequently arrested after the first few segments 
are formed, and the remainder of the process appears to be triggered when the 
larvae contact a suitable substrate. In nearly all polychaetes the larval tissues are 
resorbed with no apparent discard of material at metamorphosis. Metamorphosis 
is discussed in detail by Schroder and Hermans (1975). 

Establishment in adult habitat.—Most polychaetes appear capable of settling 
after the first few setigers have formed. However, some species can settle in 
typical larval configuration while others remain in the plankton until they have 
attained nearly adult configuration. The level of habitat selectivity appears to 
vary from species that are highly selective (Wilson 1954, 1955) to species that 
appear to follow the Thorsonian larval-rain model (Thorson 1950). After settle- 
ment all polychaetes will start feeding on food types, if not sizes, used by adults 
of the species. Species with planktotrophic larvae will at this point also shift to 
the adult food, which can be widely different from the phytoplankton consumed 
by the larvae. 

Growth phase.—Little is known about the phase of life from settlement to the 
onset of sexual maturity: At settling most species are only a fraction of their adult 
length, and densities at settlement may be entirely different from adult population 
patterns. Density regulation in the form of post-settlement dispersal or mortality 
must take place. Most species are capable of moving from one location to another, 
even the so-called sessile taxa (see Dean 1978a, b) so the growth phase may 
represent a hidden small-scale dispersal phase. 

Gamete maturation.—In most polychaetes early development of the sexual 
products takes place in gonads suspended in the coelomic cavity, with late de- 
velopment occurring while the sex cells are free in the coelom (Schroder and 
Hermans 1975). Release of sex cells from the gonads may take place as a single 
event, or be spread out over a longer period of time. Parallel to the late phases 
of gamete maturation, changes in adult morphology may also signal the onset of 
sexual maturity. Such changes are usually most drastic in semelparous species, 
but cyclic changes may also take place in interoparous forms. 

Spawning.—Polychaetes may be semelparous or iteroparous. Specimens of 
iteroparous species may spawn repeatedly within a single spawning season, or 
may spawn only once within a single season. Most polychaete species appear to 
have protracted spawning periods, lasting for several weeks to months; a few 
species are known to have extremely limited, synchronized spawning, such as 
the palolo worm, Palola viridis (e.g., Hofmann 1974). While the spawning pat- 
terns of several swarming polychaetes have been well publicized, most species 


164 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


do not swarm, but spawn in or near the adult habitat. A number of tubicolous 
forms spawn within their tubes and burrowing forms may form temporary spawn- 
ing burrows, leaving their spawn there. 

Senescence and death.—Virtually nothing is known about typical causes of 
death in iteroparous polychaetes. Most semelparous forms may die upon spawn- 
ing, and in the swarming forms tissues of several major organ-systems have been 
reported to be resorbed before the spawning takes place (Schroder and Hermans 
1975). Certain semelparous forms, such as some spionids, appear to survive the 
spawning and remain in their tubes with the developing embryos for some period 
of time. 


Material and Methods 


The material available for study is extremely limited, in that most of the dif- 
ferent kinds of information indicated by the listing of life diagram events must be 
available and studied in a single population of a species. 

Hannan et al. (1977) in a study of life histories of benthic invertebrates of 
Monterey Bay included information on larval development, the size of the repro- 
ducing females and the total number of eggs present. Species studied included 
Ameana occidentalis, Lumbrineris luti, Magelona sacculata, Mediomastus cal- 
iforniensis, Nothria elegans, Prionospio cirrifera and P. pygmaea. 

Emerson (1975) studied the population ecology of Diopatra ornata at Santa 
Catalina Island off southern California and included sufficient measurements to 
allow calculation of the necessary parameters. Beckwitt (1979) in a study of the 
population ecology of spirorbid polychaetes from southern California did a series 
of experiments on settlement and population structure allowing the inclusion of 
data for Janua brasiliensis and Pileolaria pseudomilitaris. 

Information on the reproductive biology of onuphid polychaetes has been gath- 
ered in the vicinity of the Smithsonian Marine Station at Link Port, Florida, at 
Bermuda, and in Belize; the following species have been studied in some detail: 
Kinbergonuphis simoni from Florida, Mooreonuphis jonesi at Bermuda, and K. 
pulchra in Belize. Some information is available for additional species from Flor- 
ida, and this material is used to make some testable predictions about missing 
data. 

For each species the following information was recorded: |. Size of the repro- 
ducing female (numbers of setigers as well as length and width measurements). 
2. Average egg-size. 3. Numbers of eggs produced by a female in a single repro- 
ductive event. 4. Number and distribution of reproductive events in the life of a 
single individual. 5. Estimated length of life for a single individual. 

Data are presented in tables and illustrations and compared to information 
otherwise available in the literature. 

The following biological notes indicate the basic habitat and general geograph- 
ical dispersal of principal species treated in this paper. Table | reviews some 
basic life diagram parameters for these species. 

Amaeana occidentalis Hartman (1942) (family Terebellidae) lives in temporary 
burrows in sandy and muddy environments from intertidal zones to the edge of 
the continental shelf or a little beyond; it is known from southern and central 
California, but may also be present along the whole northwest Pacific coast. 


VOLUME 96, NUMBER 1 165 


Identification of species in the genus is problematic and the exact distribution is 
currently difficult to assess. 

Diopatra ornata Moore (1910) (family Onuphidae) is tubicolous and lives at 
shelf and upper slope depths in sandy and muddy environment in the eastern 
Pacific Ocean. Emerson (1975) did a large-scale study of a single population of 
D. ornata from Santa Catalina Island. 

Janua brasiliensis (Grube, 1871) (family Spirorbidae) is a small, spirally coiled 
worm living in a calcareous tube. It is widely dispersed in warm waters in very 
shallow subtidal and intertidal areas. Beckwitt (1979) examined it as part of a 
study of intertidal spirorbids from southern California. 

Kinbergonuphis pulchra (Fauchald, 1981) (family Onuphidae) lives in relatively 
poorly constructed tubes in shallow subtidal areas. It is currently known only 
from sandy areas inside the outer edge of the barrier reef off Belize, Central 
America. 

Kinbergonuphis simoni (Santos, Day, and Rice, 1981) (family Onuphidae) lives 
in poorly constructed tubes in shallow subtidal and intertidal areas in sand. It is 
known from both sides of peninsular Florida, and is common in bays and inlets. 

Lumbrineris luti Berkeley and Berkeley (1945) (family Lumbrineridae) lives in 
sandy and muddy areas at shelf depths along the eastern Pacific Ocean. It does 
not build tubes but burrows through the substrate. 

Magelona sacculata Hartman (1961) (family Magelonidae) lives in sandy sub- 
strates at shelf depths off southern and central California. It does not build a 
permanent tube or burrow, but appears to move through the substrate more or 
less continuously. 

Mediomastus californiensis Hartman (1944) (family Capitellidae) was originally 
described from California in shallow muddy environments, but has since been 
reported widely from both coasts of the Americas. It builds semi-permanent bur- 
rows and appears tolerant of considerable environmental abuse in that it fre- 
quently is present in harbors and similar high-organic environments (Hannan et 
al. 1977). 

Mooreonuphis jonesi Fauchald (1982) (family Onuphidae) is the only onuphid 
known from shallow water in Bermuda. It is common in sand and coral debris 
and builds tubes which are covered externally with coral and shell fragments. 

Nothria elegans (Johnson, 1897) (family Onuphidae) is known from shallow 
subtidal to deep shelf depths off the west coast of the United States and Canada; 
it is tubicolous and especially common in relatively clean sandy environments. 

Pileolaria pseudomilitaris (Thiriot-Quievreux, 1965) (family Spirorbidae) ap- 
pears to be widespread in warm waters in intertidal and shallow subtidal areas; 
it is small, spirally coiled and enclosed in a calcareous tube. It was studied in 
southern California by Beckwitt (1979). 

Prionospio cirrifera (Wiren, 1883) (family Spionidae) was originally described 
from shallow water in the Arctic Ocean and has since been widely reported. It 
is possible that material studied by Hannan et al. (1977) should be referred to by 
another name (Nancy Maciolek personal communication), but since all specimens 
came from a single area and were studied over a long period of time, it appears 
safe to assume that they represent a single species. Prionospio cirrifera builds a 
small tube which it is capable of leaving, and has been reported from a variety 


166 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Table 1.—Some ecological characteristics of polychaetes considered in the analysis. The columns 
are: 1. Longevity; 2. Motility pattern of adults; 3. Feeding pattern of larvae. 


1 2 3 
Ameana occidentalis annual discretely motile feeding 
Diopatra ornata perennial sessile non-feeding 
Janua brasiliensis multiannual sessile non-feeding 
Kinbergonuphis pulchra perennial sessile non-feeding 
Kinbergonuphis simoni perennial sessile non-feeding 
Lumbrineris luti perennial motile non-feeding 
Magelona sacculata annual motile feeding 
Mediomastus californiensis multiannual discretely motile non-feeding 
Mooreonuphis jonesi perennial sessile non-feeding 
Onuphis elegans perennial sessile non-feeding 
Pileolaria pseudomilitaris multiannual sessile non-feeding 
Prionospio cirrifera multiannual discretely motile feeding 
Prionospio pygmaea multiannual discretely motile feeding 


of different environments. The particular form studied here is present along the 
eastern Pacific Ocean in muddy to sandy environments at shelf depths. 

Prionospio pygmaea Hartman (1961) (family Spionidae) was originally de- 
scribed from southern California, and while the same taxonomic considerations 
apply to this as to the preceding species, it is more likely that it is currently 
referred to by its correct name. It is known from southern and central California 
in sandy and muddy environments at shelf and slope depths. Like its congener, 
it builds small, semi-permanent tubes. 


Results 


Table 2 shows the measured and calculated values for all species for which 
adequate data were available. Table 3 reviews information available for another 
eight species, for which some information may be missing or an inadequate num- 
ber of specimens has been examined. 

The lifetime reproductive effort per female varies in a systematic fashion (Fig. 
1). Perennial species, defined as those that reproduce in at least two different 
growth seasons, have consistently low lifetime investments. For all species ex- 
amined it represents less than 10% of the volume of the females and is probably 
overestimated since it was assumed that all individuals were in their first repro- 
ductive season and would not grow before the next reproductive event. This 
assumption is known to be incorrect for Diopatra ornata and Mooreonuphis 
jJonesi (Emerson 1975; Fauchald 1982), but cannot be corrected without knowl- 
edge of the exact age of all females examined. 

The two annual species examined have considerably higher lifetime reproduc- 
tive efforts, representing more than 20% of the volume of the reproducing fe- 
males. The reproductive effort is underestimated in that it has been assumed that 
females of both species reproduce only once and die after reproduction. Some 
annual species are known to partition their spawning into several shorter bursts, 
with regeneration of the coelomic oocytes between events (Schroder and Her- 
mans 1975). The available data did not make it possible to estimate whether or 
not either species shows such patterns. 


167 


VOLUME 96, NUMBER 1 


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168 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


N.e.|K.p Perennial 


WH toe aes 


Ill, [os 


P.o. P.py M.c. Pp. J.b. : 
Multiannual 


| Fs Ss ne 


;Fecundity ( 94) 
0 20 40 60 80 100 120 140 


Fig. 1. Diagram showing the relative fertility of annual, perennial and multiannual species. The 
fecundity measure is in per cent of the total volume of the female. The species are: A.o., Ameana 
occidentalis; D.o., Diopatra ornata; J.b., Janua brasiliensis; K.p., Kinbergonuphis pulchra; K.s., 
K. simoni; L.1., Lumbrineris luti; M.s., Magelona sacculata; M.c., Mediomastus californiensis; M.j., 
Mooreonuphis jonesi; N.e., Onuphis elegans; P.p., Pileolaria pseudomilitaris; P.o., Prionospio cir- 
rifera; P. py., Prionospio pygmaea. 


Multiannual species, defined as those species that go through two or more 
generations in a single growth season, show a great deal of variability. Three 
taxa, a capitellid and two spirorbids, a have extremely high lifetime reproductive 
efforts, above 75% in all three cases; two spionids show low to moderate repro- 
ductive efforts. In the case of the two spirorbids, it is known that a single indi- 
vidual may produce up to 24 batches of eggs in a lifetime (Beckwitt personal 
communication). Spirorbids live in an unstable environment, so at least some of 
these egg-batches probably remain unrealized in the life of any given female. The 
resulting reduction in reproductive effort could be considerable, but even if one 
half of the possible egg-batches were skipped, the lifetime reproductive effort 
would still be more than twice that for any annual or perennial species. It was 
assumed that the two spionids breed only once, as the most conservative estimate 
possible. If in fact each female breeds twice, the reproductive effort per female 
will increase above the level of the annual species and be close to the level of 
the two spirorbids. 

Comparing information in Tables | and 2 leads to some interesting conclusions. 
Three major life diagram patterns appear to be present. One, represented by 
annual species, shows large reproductive efforts, moderately large eggs, and 
planktotrophic larvae. The second pattern, represented by perennial species, shows 
low reproductive efforts, moderately large to large eggs and a non-planktotrophic 
development. The third pattern, represented by the multiannual forms, shows a 
(probable) high reproductive effort, relatively small eggs, and a non-planktotro- 
phic development. These patterns are indicated in Figure 2 as Pattern I and 
Patterns Ila and IIb. The two latter patterns appear related in that both include 
forms that lack planktotrophic larvae and any organized larval or adult dispersal 
phase. 

Another interesting point is that both perennial and multi-annual forms are 
relatively sedentary as adults. Onuphids, which make up the bulk of the perennial 


169 


VOLUME 96, NUMBER 1 


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170 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


PATTERN 1 PATTERN 2A PATTERN 28 


Adult 


Metamorphic 
individual 


Larva 


EGo SIZE 


Adult 


Fig. 2. Life diagram patterns for 3 selected species of polychaetes. The species are from left to 
right: Amaeana occidentalis, Kinbergonuphis simoni and Pileolaria pseudomilitaris. The patterns are 
discussed further in the text. 


species examined, are all tubicolous. These species also appear to have lost the 
pelagic dispersal stage in that they have non-feeding, benthic larvae that are 
capable of settling immediately upon release (Emerson 1975; Fauchald personal 
observations). Kinbergonuphis simoni and Mooreonuphis jonesi both brood their 
young and release them from the parental tube at the 10- (or more) setiger stage 
(Fauchald 1982 and personal observations). Beckwitt (1979) demonstrated that 


specimens of the two spirorbid species were capable of settling immediately upon 
release from the parental tube. 


VOLUME 96, NUMBER 1 171 


Discussion 


The benthopelagic life-cycles of marine invertebrates are usually discussed in 
terms of fitness of the species, expressed as the number of offspring of each 
female (cf. Caswell 1980, 1981). The reduction in length of the pelagic phase of 
the life diagram was considered an exceptional, relatively rare phenomenon by 
Caswell (1981) who demonstrated under what conditions selection for a reduced 
larval life might take place. In fact, a variety of marine invertebrates including 
prosobranch snails, all benthic peracarid crustaceans, and various polychaetes 
show a loss of the pelagic life-phase, so the pattern can hardly be considered 
exceptional. Furthermore, among the polychaetes a variety of otherwise unre- 
lated groups show a reduction in the length of larval life; presumably this loss 
has occurred independently in each of these groups. 

The life-diagram patterns of polychaetes identified above appear difficult to fit 
with the fitness theory for a variety of reasons. First, the large, perennial forms 
among the polychaetes usually show reduced motility as adults (Fauchald and 
Jumars 1979); second, these same species also show reduced motility as larvae 
and appear to lack a dispersal phase altogether. In contrast, annual forms, which 
also include a number of large species, are frequently highly motile both as larvae 
and as adults. The multiannual forms probably show the high reproductive effort 
expected by theory (the Pianka version of McArthur and Wilson’s r- and 
K-selection theory, Pianka 1970), but do not agree with that theory in that neither 
larvae nor adults have high dispersal abilities and the reproductive effort is chan- 
neled into a few highly developed larvae rather than into the numerous broadcast 
offspring predicted by the theory. 

Shields (1982) in a discussion of the maintenance of sexual reproduction, stated 
that most life-diagram theory is based on the assumption that most species have 
large, panmictic populations and that in such populations the maintenance of sex 
leads to what Williams (1975) referred to as the cost of meiosis. A highly fit 
genotype that arises through sexual reproduction will very probably be lost to 
the next generation through well documented genetic processes. According to 
Shields (1982) asexual reproduction, which would inititally keep the highly fit 
genotype intact, eventually would lead to a ratcheting of mutations, spreading 
them through the population. Shields pointed out that in relatively small popu- 
lations of related organisms, sexual reproduction will stabilize a successful ge- 
nome and will tend to weed out deleterious mutations. Thus maintenance of sex 
in these forms, while of long-term importance in evolution, is basically a conser- 
vative feature allowing a small, successful population to maintain itself while 
conditions remain reasonably stable (in ecological time) but to retain the evolu- 
tionary flexibility that sexual reproduction implies in evolutionary time. 

The explanation for the loss of a larval dispersal phase in the perennial sessile 
polychaetes appears to involve a Shieldian inbreeding: the loss of a dispersal 
phase simply is a means of maintaining a cohesive, small interbreeding population 
so that a successful genome can be maintained. The loss of larval life then does 
not involve individual fitness per se; Caswell’s fitness argument presumably works 
for relationships within each population, but the overriding feature, the mainte- 
nance of a small interbreeding population, has been the determining factor in 
losing the dispersal phase. 


172 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


The loss of dispersal in multi-annual forms may be related to the habitat these 
forms usually occupy. These forms usually occur in relatively ephemeral envi- 
ronments. Clearly, disturbed patches are distributed in a statistically predictable 
fashion. For example, soft muddy and sandy bottoms in middle-shelf environ- 
ments are usually stable, and few disturbances occur. However, one or a few 
major storm-systems will sweep over most coastlines from time to time and the 
waves from these systems will disturb bottom sediments in water deeper than 
that reached by normal wave-trains. An interrupted band of disturbed soft sedi- 
ments is thus created along many coast-lines, but it is impossible to predict ex- 
actly where the disturbance will come in a given year: we know that the distur- 
bance will come, in a statistical sense, but cannot predict exactly where nor when. 
Most of the multi-annual forms are small, and may themselves be moved around 
with the disturbed sediments. Further, if the “‘life-expectancy’’ of a disturbed 
patch is longer than the life-expectancy of specimens of the species occupying 
the patch, then it is clearly to the advantage of the occupying species to see to 
it that its offspring do not get dispersed much beyond the outlines of the patch. 
The net effect of this ecological determinant will be exactly the same as for the 
perennial species in that each actually interbreeding population will remain small, 
and specimens found close to each other have a high probability of being closely 
related as well. 

The annual species have retained large numbers of small offspring and the 
larvae may spend a long time in the plankton. It is possible that these taxa have 
very large, panmictic populations and thus fulfill the criteria for maintenance of 
sex indicated by Williams (1975); however, there may also be valid ecological 
reasons why large numbers of larvae are maintained among these taxa. Provided 
that adults die after breeding, each population of larvae that settle will have to 
settle in an environment where the presence of adults of the same species cannot 
be used as a cue in settlement. However, Wilson (1954, 1955) demonstrated that 
the larvae of Ophelia bicornis are attracted to sand where adults have been 
present, reducing the chance-settlement in this particular species. It is also pos- 
sible that taxa in this category are unspecific in their ecological requirements and 
may be presently found in extensive environments, such as sandy and muddy 
bottoms with variable grain size, organic content, and depth. Under these cir- 
cumstances, it would be to the advantage of the species to spread its larvae as 
widely as possible and thus to maintain maximum genetic flexibility. 

The result of this study points out, that while the life-diagram theory as it has 
developed probably is correct, other features must be taken into account, and 
that the extremely heavy focus of the life-diagram theory on fitness and with the 
hidden assumption of large, panmictic populations, has made the investigators 
overlook the consequences of the fact that many animals occur in small, isolated 
or partially isolated populations in which a successful genome is being maintained 
by a loss of dispersal mechanisms. The pattern developed here is not universal: 
a great number of polychaetes do in fact disperse over wide areas and do have 
larvae that live for a long period of time in the plankton (Wilson 1968; Scheltema 
1974), but a surprisingly large number of species appear to shorten the larval life, 
either by spawning into an egg-mass (lumbrinerids, maldanids, terebellids), by 
brooding (onuphids, serpulids, spionids) or even by direct development (nerei- 
dids). In these cases, I believe the best interpretation of the curtailment of long 


VOLUME 96, NUMBER 1 173 


larval life lies in the attempt at maintaining small breeding populations, rather 
than in any optimization of fitness in terms of numbers of offspring. A second 
interpretation might be that the curtailment of larval life reduces larval wastage 
if the adult environment is hard to find and patchy. Note that these two expla- 
nations may simultaneously be correct. I believe that the fitness theory as cur- 
rently conceived might have great value in explaining differences between indi- 
vidual organisms within a confined environment. 

The reduction in lifetime reproductive effort in the perennial species indicates 
that for these species, most of which are large (for polychaetes) a larger fraction 
of the energy consumed has been shifted to growth rather than to reproduction. 
This is especially striking when comparing the lifetime reproductive effort of 
similar sized onuphids which are perennial, with the annual terebellids. 

The findings of this study are consistent with the review by Schaffer and Gadgil 
(1975) of higher plants. In some respects the coincidence may be fortuitous since 
the present emphasis has been on the role of reduced larval dispersal in main- 
taining a cohesive genetic structure in the population, rather than on the selection 
for an optimal life diagram emphasized by Schaffer and Gadgil (1975). It may be 
competitively of so great importance for a population to maintain genetic cohe- 
siveness that selection for other features in the short term becomes trivial. 

As indicated by Trendall (1982) different life diagram traits may not at all be 
closely correlated to each other. The idea of a life diagram for all populations of 
species may be spurious: each population, with the constraints built into the basic 
morphology and development of the taxon, may adapt the life diagrams to the 
local conditions under which the population survives. Despite this caveat, it ap- 
pears that the polychaetes investigated so far can be fitted into the three patterns 
described above. 

Very few of the 15,000+ known species of polychaetes were taken into account 
in erecting this system of life diagram patterns. An attempt was made at having 
as many and as varied morphological forms represented as possible, but inevita- 
bly, the few species examined represent a biased sample including species for 
which data could be easily gathered or were already present in the literature. The 
detailed structure of the life diagram patterns for polychaetes may change, but 
the basic outline will probably remain similar to the one detailed above. 


Some Predictions 


For the last several years I have been following the reproductive activity of 
selected species of polychaetes in the vicinity of the Smithsonian Marine Station 
at Link Port, Florida. For some species I have gathered sufficient information to 
include them among the species listed above. However, for most of them I have 
either inadequate numbers of specimens or some pieces of information cannot be 
gathered with the techniques I am using. 

Table 3 reviews information for these taxa. Based on life-diagram patterns I 
outlined above, I believe the following statements will, when tested, describe the 
situation for these species. 

Two of the species for which I have been gathering data are onuphids, Onuphis 
eremita oculata Hartman (1951) and Diopatra cuprea (Bosc, 1802). For both 
species I lack crucial data, but the average egg-sizes for both indicate that the 


174 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


life diagram for each should be similar to that found for other onuphids, with low 
reproductive effort, small numbers of eggs and lack of pelagic larvae, or at least 
tube-brooding until larvae are competent to settle. 

Axiothella sp., a maldanid, is known to discharge eggs into an eggmass; the 
egg-size and average numbers of eggs per batch indicate a reduced larval devel- 
opment, without a feeding larval stage. The reproductive effort of the only com- 
plete female sampled indicates that the species should live for at least two sea- 
sons. 

Glycinde sp., a goniadid, has a relatively high reproductive effort combined 
with small eggs; it is suspected that this species may be annual and have a pelagic, 
planktotrophic larva. 

Streblosoma sp. of the family Terebellidae, has relatively large eggs, and mod- 
erately high reproductive effort. Related species are known to spawn into a loose- 
ly organized egg-mass, which deteriorates rapidly and releases larvae, which, 
while they may be planktotrophic, are capable of settling shortly after release 
from the egg mass. Based on the available information, it appears that Streblo- 
soma will show a similar pattern. The species may be an annual. 

The capitellids studied by Grassle and Grassle (1974, 1976) show a bewildering 
array of different life-diagrams. Some taxa have planktotrophic larvae, others 
have a reduced larval life or direct development, the average egg-diameters ap- 
pear well correlated with the developmental pattern exhibited (see also Schroder 
and Hermans 1975, and Hermans 1979). The consequences in terms of life-dia- 
grams are that the forms with a planktonic larva are capable of dispersing rapidly 
and widely, whereas the forms without such larvae are capable of building up 
larger populations once a suitable habitat has been located. The only specimen 
of Capitella measured in the current study had large eggs and thus should belong 
to the taxa with a reduced larval development. The different recruitment strate- 
gies of the capitellids may well determine longevity and thus differential resource 
utilization by each species, and may thus explain why the Grassles found more 
than a single species in a sample in certain areas off Massachusetts. 

Two specimens of Haploscoloplos fragilis measured during the current study 
show egg sizes similar to those measured for Scoloplos armiger, another orbiniid, 
by Anderson (1959). The reproductive effort is very low and very few segments 
contained eggs in both specimens. It is suggested that this orbiniid shows a mixed 
strategy: the larvae are planktotrophic, but the individual specimens live for at 
least two seasons or alternatively, are capable of spawning more than once, each 
well separated out in the life of the female. The two individuals, both complete, 
are very different in size, indicating that perhaps the latter alternative may be the 
correct one. 

A single specimen of a species of Ophelina was measured. The egg diameters 
are similar to those measured for Armandia bioculata by Hermans (1979) and it 
is predicted that the life diagram will be characteristic of an annual species, with 
high reproductive effort, small, planktotrophic larvae and relatively lengthy 
planktonic life. 

Another interesting prediction is related to the structure of the deep sea ben- 
thos. The dominating benthic forms at least among the polychaetes in deep water 
are all forms that fit with the pattern found above for multi-annual forms (see 
Hartman 1965, Hartman and Fauchald, 1971). The shallow water multi-annual 


VOLUME 96, NUMBER 1 175 


forms are associated with areas of randomly disturbed sediments or other unsta- 
ble, unpredictable environments. The dominance of species with a similar life 
diagram in the deep sea benthos indicates that a similar disturbance pattern may 
be present in the apparently stable deep sea benthos. As indicated by Dayton 
and Hessler (1972) this disturbance is very probably biological in nature. 


Acknowledgments 


This study started with a lecture I gave during the class in polychaete biology 
at Catalina Marine Science Center, University of Southern California, in 1979. 
Bill Kennedy and Fred Piltz, the assistants in that class, and also the students 
encouraged me to develop the topic further; particularly Bill Kennedy was in- 
strumental in getting me started. The reaction to a paper I gave at the annual 
meeting of American Society of Zoologists in Tampa indicated that the material 
would create a great deal of interest, if not agreement. Various versions of the 
paper have been read by Meredith L. Jones, Mary E. Rice, Peter A. Jumars, 
Kirk Fitzhugh, and David E. Russell, all of whom gave their candid opinions but 
should not be held responsible for its content. 

This is contribution #116 from the Smithsonian Marine Station at Link Port, 
Florida. 


Literature Cited 


Akesson, B. 1967. The embryology of the polychaete Eunice kobiensis.—Acta Zoologica 48:141-— 
192, 22 figs. 

Anderson, D. T. 1959. The embryology of the polychaete Scoloplos armiger.—Quarterly Journal 
of Microscopical Science 100:89-166. 

—. 1961. The development of the polychaete Haploscoloplos fragilis.—Quarterly Journal of 

Microscopical Science 102:257—272. 

1966. The comparative embryology of the Polychaeta.—Acta Zoologica 47: 1—42. 

Beckwitt, R. D. 1979. Genetic structure of Pileolaria pseudomilitaris (Polychaeta: Spirorbidae) in 
southern California.—Dissertation, University of Southern California, Los Angeles, 126 pp., 
11 figs., 13 tbs. 

Blake, J. A. 1969. Reproduction and larval development of Polydora from northern New England 
(Polychaeta: Spionidae).—Ophelia 7: 1-63. 

Caswell, H. 1980. On the equivalence of maximising reproductive value and maximising fitness.— 
Ecology 61(1):19—24, | tb. 

——. 1981. The evolution of “‘mixed”’ life histories in marine invertebrates and elsewhere.— 
American Naturalist 117(4):529-536, | fig. 

Dayton, P. K., and R. R. Hessler. 1972. Role of biological disturbance in maintaining diversity in 
the deep sea.—Deep-Sea Research 19:199-208. 

Dean, D. 1978a. Migration of the sandworm, Nereis virens, during winter nights.—Marine Biology 
45:165-173. 

——. 1978b. The swimming of bloodworms (Glycera spp.) at night, with comments on other 
species.—Marine Biology 48:99-104. 

Eckelbarger, K. J. 1980. An ultrastructural study of oogenesis in Streblospio benedicti (Spionidae), 
with remarks on diversity of vitellogenic mechanisms in Polychaeta.—Zoomorphologie 94:24 1— 
263, 32 figs. 

Emerson, R. R. 1975. The biology of a population of Diopatra ornata at Santa Catalina Island, 
California.—Dissertation, University of Southern California, Los Angeles, 325 pp., 30 figs., 27 
tbs. 

Fauchald, K. 1982. Description of Mooreonuphis jonesi, a new species of onuphid polychaete from 
shallow water in Bermuda, with comments on variability and population ecology.—Proceed- 
ings of the Biological Society of Washington, 94(4):807-825. 


176 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


, and P. A. Jumars. 1979. The diet of worms: a study of polychaete feeding guilds.—Ocean- 

ography and Marine Biology, An Annual Review 17:193—284, 23 figs., 33 tbs. 

Grassle, J. F., and J. P. Grassle. 1974. Opportunistic life histories and*genetic systems in marine 
benthic polychaetes.—Journal of Marine Research 32(2):253—284, 11 figs., 2 tbs. 

Grassle, J. P., and J. F. Grassle. 1976. Sibling species in the marine pollution indicator Capitella 
(Polychaeta).—Science 192(4239):567-569. 

Hannan, C. A., L. W. Hulberg, K. M. Mawn, and J. W. Nybakken. 1977. A study to develop 
standard procedures for life history analysis of benthic invertebrates for biological monitoring 
in marine and estuarine environments.—Moss Landing Marine Laboratories, California State 
University Consortium, Moss Landing, California, 217 pp. 

Hannerz, L. 1956. Larval development of the polychaete families Spionidae Sars, Disomidae Mesnil 
and Poecilochaetidae n. fam. in the Gullmar Fjord (Sweden).—Zoologiska Bidrag fran Uppsala 
31:1-204, 57 figs. 

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

other North Atlantic areas.—Allan Hancock Foundation Publications, Occasional Paper 28:1— 

378, 52 pls. 

, and K. Fauchald. 1971. Deep-water benthic polychaetous annelids off New England to 

Bermuda and other North Atlantic areas Part IJ.—Allan Hancock Monographs in Marine Bi- 

ology 6:1—327, 34 pls. 

Hermans, C. O. 1979. Polychaete egg sizes, life histories and phylogeny. In S. E. Stancyk, ed. 
Reproductive ecology of marine invertebrates.—The Belle W. Baruch Library in Marine Sci- 
ence 9:1—9, | fig., 3 tbs. 

Hofmann, D. K. 1974. Maturation, epitoky and regeneration in the polychaete Eunice siciliensis 
under field and laboratory conditions.—Marine Biology 25:149-161. 

Knight-Jones, E. W., P. Knight-Jones, and P. J. Vine. 1972. Anchorage of embryos in Spirorbinae 
(Polychaeta).—Marine Biology 12:289-294, 2 figs. 

Mauro, N. A. 1975. The premetamorphic developmental rate of Phragmatopoma lapidosa Kinberg 
1867, compared with that in temperate sabellariids (Polychaeta, Sabellariidae).—Bulletin of 
Marine Science 25(3):387-392. 

Menge, B. A. 1975. Brood or broadcast? The adaptive significance of different reproductive strat- 
egies in the two intertidal sea stars Leptasterias hexactis and Pisaster ochraceus.—Marine 
Biology 31:87—100. 

Palmer, A. R., and R. R. Strathmann. 1981. Scale of dispersal in varying environments and its 
implication for life histories of marine invertebrates.—Oecologia 48:308—318, 8 figs., 2 tbs. 

Pianka, E. On r- and K-selection.—American Naturalist 104:592—597. 

Pielou, E. C. 1981. The usefulness of ecological models: a stock-taking.—Quarterly Review of 
Biology 56:17-31. 

Schaffer, W. M., and M. D. Gadgil. 1975. Selection for Optimal life histories in plants. Pp. 142— 
157 in M. L. Cody and J. M. Diamond (eds.). Ecology and evolution of communities.—The 
Belknap Press of Harvard University Press. 5 figs., 2 tbs. 

Scheltema, R. S. 1974. Relationship of dispersal to geographical distribution and morphological 
variation in the polychaete family Chaetopteridae.—Thalassia Jugoslavica 10(1/2):297-312. 

Schroder, P. C., and C. O. Hermans. 1975. Annelida: Polychaeta. Jn A. C. Giese and J. S. Pearse 
(eds.). Reproduction of marine Invertebrates. III, Annelids and Echiurans, Chapter 1:1—213. 

Shields, W. M. 1982. Inbreeding and the paradox of sex: a resolution?—Evolutionary Theory 5(5): 
245-279, 4 figs., 5 tbs. : 

Smith, R. I. 1950. Embryonic development in the viviparous polychaete Neanthes lighti.—Journal 
of Morphology 87:417—466. 

Stearns, S.C. 1976. Life-history tactics: a review of ideas.—Quarterly Review of Biology 51:3—47. 

Thorson, G. 1946. Reproduction and larval development of Danish marine bottom invertebrates, 
with special reference to the planktonic larvae in the Sound (Oresund).—Meddelelser fra Kom- 
missionen for Danmarks Fiskeri- og Havundersogelser, serie plankton 4(1):1—523, 199 figs. 

——. 1950. Reproductive and larval ecology of marine bottom invertebrates.—Biological Re- 
views, Cambridge 25:1-45. 

Trendall, J. T. 1982. Covariation of life history traits in the mosquitofish, Gambusia affinis.— 
American Naturalist 119(6):774-783, 4 tbs. 

Wiley, E.O., and D. R. Brooks. 1982. Victims of history—a nonequilibrium approach to evolu- 

tion.—Systematic Zoology 31(1):1—24, 7 figs., 2 tbs. 


VOLUME 96, NUMBER 1 Gia 


Williams, G. C. 1975. Sex and evolution.—Princeton University Press Monographs in Population 
Biology 8:1—200, 17 figs. 

Wilson, D. P. 1954. The attractive factor in the settlement of Ophelia bicornis Savigny.—Journal 
of the Marine Biological Association of the United Kingdom 34:361-380. 

———. 1955. The role of micro-organisms in the settlement of Ophelia bicornis Savigny.—Journal 
of the Marine Biological Association of the United Kingdom 34:531—543. 

——. 1968. The settlement behaviour of the larvae of Sabellaria alveolata (L.)—Journal of the 
Marine Biological Association of the United Kingdom 48:387-435. 


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


PROC. BIOL. SOC. WASH. 
96(1), 1983, pp. 178-180 


ARGYROTHECA ARGUTA, A NEW SPECIES OF 
BRACHIOPOD FROM THE MARSHALL 
ISLANDS, WESTERN PACIFIC 


Richard E. Grant 


Abstract.—Argyrotheca arguta is characterized by small size (maximum width 
2.8 mm), white, clear or translucent shell, and weak to absent costae. It occupies 
cryptic habitats under coral fronds or in recesses, and has been found on the sea 
side and the lagoon side of the reefs at Enewetak and Bikini atolls. 


A book on the natural history of Enewetak is in preparation under the auspices 
of the Mid Pacific Research Laboratory at Enewetak. I was invited to write up the 
brachiopods, based upon my observations of them there and the collections in the 
National Museum. Only three species have been found at Enewetak, yet despite all 
the work that has been done, there remains unnamed a species of Argyrotheca. 
This has been a mild inconvenience for years (e.g., Cooper 1954) but previously the 
collection was too small to allow a confident decision as to whether these small 
shells represent a new species or merely babies of a described form. Now enough 
material is at hand to reveal features other than mere size that indicate that the 
Enewetak form is a distinct species, apparently endemic to the Marshall Islands. 

The editors of the book on Enewetak wish to avoid the introduction of new 
taxa, hence this short note to describe Argyrotheca arguta, new species. 


Phylum Brachiopoda Dumeril 
Class Articulata Huxley 
Order Terebratulida Waagen 
Superfamily Terebratellacea King 
Family Megathyrididae Dall 
Genus Argyrotheca Dall 
Argyrotheca arguta, new species 
Fig. | 


Diagnosis.—Shell small; known width less than 3 mm; outline heart-shaped 
with slightly emarginate anterior margin; profile biconvex with inflated dorsal 
valve and rather flat ventral valve; color white to light tan or clear, translucent; 
some shells with low and indistinct costae, most lacking costae. 

Ventral valve with open triangular pedicle foramen flanked by strong divergent 
hinge teeth extending nearly to anterior margin. Dorsal valve with deep hinge 
sockets formed between valve edge and socket plates; loop short, simple, with 2 
descending branches circling laterally and converging on median septum; some 
adults with serrate anterior margins. 

Holotype.—USNM 265875: the largest shell in the collection with width 2.8 mm. 

Etymology.—From the Latin ‘‘argutus’’ = “‘clear,’’ referring to the translucent 
Shell. 

Comparison.—This species is characterized by small size and translucent shell 


VOLUME 96, NUMBER 1 179 


Fig. 1. Argyrotheca arguta: 11 shells attached to underside of coral frond from USNM loc. 32014 
on the lagoon side of Igurin Island, Enewetak Atoll, depth 14 m. The long arrow points to the holotype, 
the short arrows to other specimens (x4). 


that lacks the reddish color or stripes that typify other species of Argyrotheca, 
and lacks the radial costae that most species show. It was recognized long ago 
that these shells did not fit easily into any of the described species (Cooper 1954) 
but they were not described for fear that they were merely juveniles. The present 
sample is larger than any studied previously and it contains very small shells as 
well as a majority that seem to have ceased growth at a width between 2 and 3 
mm. The species is currently known only from Enewetak and Bikini, but further 
exploration probably will reveal it throughout the Marshall Islands. It is associ- 
ated with the small cemented brachiopod Thecidellina congregata Cooper at both 
atolls. 

Species of Argyrotheca in the Caribbean Sea are larger than A. arguta and 
have costae and red stripes. Two species in the Pacific, A. australis and A. mayi 
from Australia and Tasmania, also differ from A. arguta by larger size, strong 
costation, and colored radial stripes. 

Habitat.—Argyrotheca arguta was found in two abundant patches at Enewe- 
tak, both on the ceilings of recesses in the reef. The samples were obtained by 
divers who observed the habitat. The sample at Bikini was obtained by dredge, 
so the habitat was not seen. One Enewetak sample is from the sea side of the 
reef at Rigili Island at a depth of about 30 m; the other is from the lagoon side 
of Igurin at 14 m. The dredged sample from Bikini was taken from about 100 m 
depth on the south side of the atoll (Cooper 1954). 


180 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Literature Cited 


Cooper, G. A. 1954. Recent brachiopods. Bikini and nearby atolls, Part 2. Oceanography (biolog- 
ic) —U.S. Geological Survey Professional Paper 260-G:315-318, pl. 80-81. 


Department of Paleobiology, Smithsonian Institution, Washington, D.C. 20560. 


PROC. BIOL. SOC. WASH. 
96(1), 1983, pp. 181-188 


HARBANSUS SLATTERYI, A NEW SPECIES OF 
MYODOCOPINE OSTRACODE FROM THE 
GREAT BARRIER REEF OF AUSTRALIA 

(PHILOMEDIDAE) 


Louis S. Kornicker 


Abstract.—Harbansus slatteryi, new species, is described and illustrated. The 
specimens were collected from the Lizard Island Group, Great Barrier Reef, 
Australia. The genus had not previously been recorded from Australia. 


The genus Harbansus Kornicker, 1978, was proposed for numerous species 
collected in the Atlantic Ocean, and also in the Pacific Ocean as far west as 
Hawaii. The present report extends the range farther west and south to the Great 
Barrier Reef in the Coral Sea off Queensland, Australia. 


Harbansus Kornicker, 1978 


Type-species.—Harbansus bradmyersi Kornicker, 1978. 


Harbansus slatteryi, new species 
Figs. 1-4 


Etymology.—The species is named for Peter N. Slattery who collected some 
of the specimens. 

Material.—Palfrey Island, Lizard Island Group, Great Barrier Reef, Queens- 
land, Australia. Collected by Peter N. Slattery, Sept-Oct 1977: core sample, 
water depth 12 m, quartz sand substrate, | adult female, USNM 157790, holotype; 
station 8, at far edge of reef flat adjacent to coral head at edge of coral reef, 
probably exposed only twice a year at extreme low tides, fine to medium fine 
sand, 5 large cores (each 0.0176 m?) sieved through 500 « mesh, | adult female, 
USNM 158490, paratype. West of Lizard Island Research Station, off Casuarina 
Beach, station AC-4A, 27 May 1980, 200-300 yards offshore of sandy beach, 
small sand patch between coral heads in patch reef near edge, about 2 m deep, 
3 adult males, USNM 158491, paratypes, collected by Anne C. Cohen. Lagoon 
south of Lizard Island, between Lizard Island and Palfrey Island, station AC-6, 
28 May 1980, off sandy beach with | palm tree, about 200-300 yards from shore, 
sparse Thalassia bed in very silty sand, about 2 m deep, collected by Anne C. 
Cohen, paratypes: | adult male, USNM 158492; 1 adult male, USNM 158627; | 
adult female, USNM 159092; | adult male, USNM 159093. 

Description of adult female (Figs. 1-3).—Carapace elongate with long narrow 
rostrum and caudal process (Figs. |, 2). Ornamentation (Fig. 1): Each valve with 
3 horizontal ribs and additional smaller rib near and parallel to anterodorsal mar- 
gin; On some specimens, anterior end of rib dorsal to adductor muscle attachment 
area joining anterior end of anterodorsal rib to form small process extending very 
Slightly past anterior end of rostrum (Fig. 1); on other specimens, anterodorsal 


182 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 1. Harbanus slatteryi, adult female, holotype, USNM 157790, lateral view of complete spec- 
imen, length 1.00 mm. 


rib not reaching anterior margin; midrib passing through area of central adductor 
muscle attachments; its anterior end bending ventrally to join with anterior end 
of ventral rib to form small process extending slightly past anteroventral margin 
of valve; low vertical rib connecting anterior ends of rib dorsal to adductor muscle 
attachments and midrib. Surface of valves with abundant oval fossae (Fig. 1). 
Long bristles forming row near distal end of caudal process (Fig. 2c), along ventral 
margin of each valve, and sparsely distributed over valve surface (these bristles 
not shown in illustrations). 

Infold (Fig. 2): Infold of rostrum with 4 long bristles forming row (Fig. 2a, b); 
2 bristles present at inner end of incisure; anteroventral infold with | short bristle 
ventral to inner end of incisure and with ridges paralleling valve margin; inner 
edge of infold anterior to caudal process with about 6 small bristles; ridge forming 
anterior end of pocket in caudal process with 6 frondlike bristles forming row 
(Fig. 2c); 2 minute bristles along posterior edge of caudal process. 

Selvage (Fig. 2b): Narrow lamella with short marginal fringe present along 
dorsal and anterior edges of rostrum (Fig. 2b) and becoming wider along ventral 
margin of rostrum; narrow lamella with short marginal fringe present along an- 
teroventral edge of valve becoming wider in vicinity of incisure; ventral lamella 
wide with long marginal fringe; narrow lamella with short fringe present along 
posterior margin of valve; lamella apparently absent along posterior edge of cau- 
dal process. 

Size: USNM 157790, length 1.00 mm, height 0.53 mm; USNM 158490, length 
1.07 mm, height 0.60 mm; USNM 159092, length 1.08 mm, height 0.57 mm. 


VOLUME 96, NUMBER | 183 


Fig. 2. Harbanus slatteryi, adult female, holotype, USNM 157790: a, Anterior of right valve, 
inside view; b, Rostrum and incisure of left valve, including selvage, inside view; c, Caudal process 
of right valve, inside view. 


First antenna (Fig. 3a): Joints 1-4 with surface spines forming rows. 2nd joint 
with | dorsal bristle. 3rd joint short, with 3 bristles, | ventral, 2 dorsal. 4th joint 
elongate, with terminal bristles, 2 ventral, 1 dorsal. 5th joint elongate; sensory 
bristle with 2 short proximal filaments, 3 longer distal filaments, and 2 minute 
spines at tip. 6th joint fused with 5th, with | small medial bristle. 7th joint: a- 
bristle about twice length of bristle of 6th joint; b-bristle about twice length of a- 
bristle, shorter than sensory bristle of Sth joint, with | distal marginal filament, 
and tip with 2 minute spines; c-bristle with | short proximal filament and | short 
and 2 long distal filaments. Eighth joint: d- and e-bristles about same length as 
sensory bristle, bare with blunt tips; f- and g-bristles about same length as sensory 
bristle; f-bristle with 1 short proximal filament and 3 longer distal filaments, and 
with 2 minute spines at tip; g-bristle with | short proximal filament and 2 longer 
distal filaments, and with 2 minute spines at tip; all filaments on bristles of joints 
5, 7, 8 with spine at tip. 

Second antenna (Fig. 3b): Protopodite with few faint spines forming rows on 
medial and lateral surfaces near dorsal margin. Endopodite 2-jointed: Ist joint 
with 2 short proximal anterior bristles; 2nd joint elongate with bare rounded or 
tapering tip and | long spinous ventral bristle. Exopodite: Elongate Ist joint with 
1 minute terminal medial bristle; bristle of 2nd joint reaching past 9th joint, with 
about 10 stout ventral spines followed by numerous slender ventral spines; bris- 
tles of joints 3—7 with few long proximal dorsal hairs; bristles of joints 3—8 with 
stout proximal ventral spines followed by long natatory hairs on both margins; 
9th joint with minute lateral tooth and 3 bristles: 1 long with stout proximal ventral 
spines and distal natatory hairs, | medium with short marginal spines, | short 


184 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


and either bare or with few faint spines; joints 3-8 with basal spines (spines 
slightly larger on some distal joints); basal spine of 8th joint about one-half length 
of 9th joint; joints 2-8 with minute spines forming row along distal margin. 

Mandible (Fig. 3c): Coxale endite hirsute and spinous, without small bristle 
near base. Basale: dorsal margin with 3 bristles (1 near middle, 2 subterminal; 
ventral margin with | terminal bristle with short marginal spines; medial surface 
spinous, with 4 bristles near ventral margin (3 proximal, | near middle) (2 dashed 
bristles broken off illustrated limb but present on right limb); lateral surface with 
2 bristles with long spines near ventral margin. Exopodite about three-fourths 
length of dorsal margin of Ist endopodial joint, with 2 subterminal bristles (lateral 
and proximal of these much longer than other and spinous). Ist endopodial joint 
with medial spines and 3 spinous ventral bristles. 2nd endopodial joint: dorsal 
margin with proximal spines and 6 bristles near middle forming 2 groups (1 bristle 
in proximal group, 5 in distal group); ventral margin with 4 bristles forming 2 
distal groups (2 bristles in each group); medial surface with spines forming rows. 
End joint with 2 long claws with faint ventral spines, and 3 slender bristles. 

Maxilla (Fig. 3d, e): Precoxale and coxale with dorsal fringe of long hairs. 
Dorsal margin of coxale with hirsute dorsal bristle. Endite I wider than endites 
II and III, with 6 bristles; endite II also with 6 bristles; endite III with about 7 
distal bristles and | proximal lateral bristle. Basale with 3 bristles along distal 
margin. Exopodite small, with 3 bristles (2 long, | short) (Fig. 3e). Endopodite: 
Ist joint with | spinous alpha-bristle and 2 beta-bristles (beta-bristles obscure and 
number approximate); end joint with about 7 claws and bristles. 

Fifth limb (Fig. 3f-h): Endite I with 1 bristle; endite II with about 5 bristles; 
endite III with 6 or more bristles. Exopodite: Ist joint with main tooth consisting 
of 3 bilobed teeth and 2 pointed smooth peglike teeth, | at each end of the bilobed 
teeth (Fig. 3f, g); 1 small spinous bristle proximal to inner peglike tooth; additional 
bristles on Ist joint obscure; large tooth of 2nd joint triangular with large node 
near middle of inner margin (Fig. 3h); bristles of 2nd joint obscure; 3rd joint with 
2 bristles on outer lobe and 3 on inner lobe; 4th and 5th joints fused, hirsute, with 
total of 6 bristles. 

Sixth limb (Fig. 3i): | short spinous bristle in place of epipodial appendage; 
endite I small, with 2 short spinous bristles; endite I] narrow, with 3 spinous 
terminal bristles; endite III broad with 6 spinous terminal bristles; endite IV broad 
with 5 spinous terminal bristles; end joint not prolonged posteriorly, with 5 or 6 
bristles (3 or 4 anterior bristles spinous, 2 posterior bristles hirsute); limb hirsute. 

Seventh limb (Fig. 3j): Each limb with 6 bristles (2 proximal, 4 terminal), each 
bristle with marginal spines and 1-4 bells; terminal comb with about 5 recurved 
teeth; 4 or 5 short pegs present opposite comb. 

Furca (Fig. 3k): Each lamella with 6 claws; claws 1, 2, 4 stout, claw 3 and 
claws following claw 4 slender. Claw | with teeth along posterior edge and also 
with fairly large medial teeth forming row; distal medial tooth especially promi- 
nent; remaining main claws with teeth along posterior edge; edge of lamella fol- 
lowing claw 6 hirsute. 

Bellonci organ (Fig. 31): Elongate with rounded tip and 2 or 3 weak proximal 
sutures. 

Eyes (Fig. 31): Medial eye with light brown pigment. Lateral eye small, unpig- 
mented, with 2 light-amber ommatidia. 


VOLUME 96, NUMBER | 


185 


Ss. 
eee 


Hi? 


Ly 
WES 
Sp 
SY 
Ly 
oy 


= =m Scena, . 
seal 


— srw 
LT TTF 


Fig. 3. 


Harbanus slatteryi, adult female, holotype, USNM 157790: a, Left Ist antenna, medial 
view; b, Endopodite and distal part of protopodite of left 2nd antenna, lateral view; c, Left mandible, 


medial view; d, Right maxilla, medial view; e, Exopodite of right maxilla, medial view as seen through 
limb shown in “‘d’’; f, Distal part of left fifth limb, anterior view; g, Detail of tooth of Ist expodal 


joint of Sth limb shown in “‘f’; h, Distal part of right 5th limb, tooth of Ist exopodal joint not shown 
(posterior view); i, 6th limb; j, 7th limb; k, Posterior of body showing right lamella of furca, right 
genitalia, right Y-sclerite, right segment of girdle; 1, Anterior of body showing left lateral eye, medial 
eye and bellonci organ, and upper lip. 


Upper lip (Fig. 31): Consisting of single lobe. 


with attached spermatophore. 


Genitalia (Fig. 3k): Consisting of sclerotized ring on each side of body, each 
Y-sclerite (Fig. 3k): Typical for genus. 


Eggs: USNM 157790 with 3 eggs in marsupium; USNM 158490 with 2 eggs. 
Description of adult male (Fig. 4).—Except for having a slightly broader ros- 


186 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


—— — 


\ 
| 
| 


SS 


Fig. 4. Harbanus slatteryi, a, b, adult male paratype, USNM 158492, length 0.95 mm; a, Lateral 
view of complete specimen showing left valve with outline of ribs and 11 typical fossae, position of 
central adductor muscle attachments (dashed circle), position of left lateral eye (stippled), left lamella 
of furca, and left copulatory organ; b, Anterior of body showing left lateral eye with few ommatidia 
along dorsal margin, medial eye and bellonci organ, and upper lip. c+, Adult male paratype, USNM 
158627, length 0.97 mm: c, Left Ist antenna, medial view; d, Endopodite, distal part of protopodite, 
Ist joint of exopodite of left 2nd antenna, medial view; e, Left mandible, medial view; f, Maxilla; g, 
Distal part of 5th limb; h, 6th limb; i, 7th limb. 


trum and caudal process, carapace of male similar in shape and ornamentation 
to that of adult female (Fig. 4a). 

Infold: Infold of rostrum with 4 long bristles; 2 bristles present at inner end of 
incisure; anteroventral infold with | short bristle ventral to inner end of incisure 
and with ridges paralleling valve margin; inner edge of infold anterior to caudal 
process with about 4 small bristles; ridge forming anterior edge of pocket in caudal 
process with 5 or 6 frondlike bristles forming row; 2 or 3 minute bristles forming 
row along posterior edge of caudal process. 

Selvage: Similar to that of female. 


VOLUME 96, NUMBER 1 187 


Size: USNM 158492, length 0.95 mm, height 0.46 mm; USNM 158491, 3 spec- 
imens, length 0.98 mm, height 0.48 mm, length 0.94 mm, height 0.45 mm, length 
0.95 mm, height 0.48 mm; USNM 158627, length 0.97 mm, height 0.47 mm. 

First antenna (Fig. 4c): Ist joint with medial spines forming rows. 2nd joint 
with few lateral spines and | dorsal bristle with long marginal hairs. 3rd joint 
short with small medial spines and 3 bristles (2 dorsal, 1 medial near ventral 
margin); 4th joint with | bare dorsal terminal bristle. Sth joint small, wedged 
ventrally between 4th and 6th joints; sensory bristle with bulbous proximal part 
with abundant filaments (filaments not shown on illustrated limb), and stem with 
3 filaments near middle and 2 minute spines at tip (only proximal part of stem 
shown on illustrated limb). 6th joint with bare medial bristle. 7th joint: a-bristle 
spinous, longer than bristle of 6th joint; b-bristle about twice length of a-bristle, 
with 2 filaments near middle and bifurcate tip; c-bristle longer than sensory bristle 
of 5th joint, with 7 or 8 marginal filaments and bifurcate tip. 8th joint: d- and e- 
bristles slightly shorter than c-bristle, bare with blunt tips (bristles not shown on 
illustrated limb); f-bristle slightly shorter than c-bristle, with 6 marginal filaments 
and bifurcate tip; g-bristle same length as c-bristle, with 5 marginal filaments and 
2 spines at tip (bristle not shown on illustrated limb). 

Second antenna (Fig. 4d): Protopodite bare. Endopodite 3-jointed: Ist joint 
short with 4 short anterior bristles; 2nd joint elongate with 2 long proximal bris- 
tles; 3rd joint elongate, reflexed, with 2 small bristles near sclerotized beaklike 
tip, and minute processes along inner margin. Exopodite: Ist joint elongate with 
minute terminal medial bristle; 2nd joint slightly longer than 3rd joint; bristle of 
2nd joint with 3 or 4 long proximal hairs on dorsal margin and about 6 proximal 
hairs on ventral margin; hairs on ventral margin of bristle followed by 3 stout 
spines; both margins of bristle distal to spines with natatory hairs; bristles of 
joints 3-8 longer than bristle of 2nd joint, with natatory hairs but no spines; 9th 
joint with 3 bristles (1 long, stout, with natatory hairs; | medium with few ventral 
spines in addition to natatory hairs; | short with natatory hairs); joints 2-8 with 
slender spines forming row along distal margin; basal spine observed on 8th joint 
only. 

Mandible (Fig. 4e): Coxale endite represented by faint minute spine. Basale: 
medial surface with 4 short bristles near ventral margin (3 proximal, | near mid- 
dle); ventral margin with 3 long spinous bristles (2 proximal of these with bases 
on lateral surface slightly inward from ventral margin); dorsal margin with 3 long 
bristles, 1 near middle, 2 terminal. Exopodite spinous, reaching past middle of 
dorsal margin of Ist endopodial joint, with 1 long and | short bristle. 1st endo- 
podial joint with few medial spines and 3 ventral bristles (2 long, | short). 2nd 
endopodial joint: medial surface with spines forming rows; ventral margin with 
bristles forming 2 distal groups, each with 2 bristles: middle of dorsal margin with 
6 bristles. End joint with 2 stout claws and 3 bristles (1 dorsal, lateral: 2 ventral). 

Maxilla (Fig. 4f): Limb reduced and with weakly developed bristles. 

Fifth limb (Fig. 4g): Epipodial appendage with 35 bristles. Endites weakly de- 
veloped with few bristles. Exopodite: 3rd joint with 2 ringed outer bristles, inner 
bristles obscure; remaining joints obscure, with total of about 5 bristles. 

Sixth limb (Fig. 4h): A single spinous bristle in place of epipodial bristle. Endite 
I with 3 bristles; endite 2 with 3 terminal bristles; endite III with 5 or 6 bristles; 


188 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


endite IV with 4 or 5 bristles. End joint not prolonged posteriorly, with 6 or 7 
bristles (4 or 5 anterior bristles either spinous, hirsute, or with long proximal hairs 
and short distal spines; 2 posterior bristles stout, hirsute); limb hirsute. 

Seventh limb (Fig. 41): 4 bristles in terminal group, 2 on each side, each with 
4 bells; some bristles with few faint, distal, marginal spines; proximal bristles 
absent. Terminus with comb of 3 teeth, each lateral tooth with long proximal 
spines; surface opposite comb without pegs. 

Furca (Fig. 4a): Similar to that of female except claw | without stout, distal, 
medial tooth present on female. 

Bellonci organ (Fig. 4b): Elongate with round tip and about 4 proximal sutures. 

Eyes (Fig. 4b): Medial eye bare with reddish brown and black pigment. Lateral 
eye about same size as medial eye or slightly larger, with about 7 ommatidia but 
difficult to count because of black pigment. 

Upper lip (Fig. 4b): Consisting of single lobe. 

Genitalia (Fig. 4a): Each copulatory limb elongate, lobate, with terminal pro- 
cess with marginal teeth. 

Y-sclerite: Similar to that of female. 

Comparisons.—Previously described species of Harbansus having lateral ribs 
on the carapace are H. paucichelatus (Kornicker, 1958:233), H. bradmyersi Kor- 
nicker (1978:24), and Harbansus species B (Kornicker, 1978:49). The adult fe- 
males of the last two species differ from the new species H. slatteryi in not having 
natatory hairs on the exopodal bristles of the 2nd antenna. The adult female 
mandible of H. slatteryi differs from all previously described species of the genus 
in having only two claws instead of three on the end joint of the mandible. The 
adult female of H. slatteryi bears four or five (usually five) pegs on the tip of the 
7th limb compared to only two or three pegs on H. paucichelatus. 


Acknowledgments 


My thanks to the following individuals for their help: Peter N. Slattery, Moss 
Marine Laboratory, Moss Island, California, and Anne C. Cohen, Department of 
Invertebrate Zoology, Smithsonian Institution, for collecting specimens studied 
herein; to Carolyn Bartlett Gast, Smithsonian Institution, for the shaded rendering 
of the carapace, and to Thomas E. Bowman, Smithsonian Institution, for com- 
menting on the manuscript. 


Literature Cited 


Kornicker, Louis S. 1958. Ecology and taxonomy of Recent marine ostracodes in the Bimini area, 
Great Bahama Bank.—Publications of the Institute of Marine Science (The University of Tex- 
as) 5:194—300. 

——. 1978. Harbansus, a new genus of marine Ostracoda, and a revision of the Philomedidae 
(Myodocopina).—Smithsonian Contributions to Zoology 270, 75 pp. 


Department of Invertebrate Zoology, Smithsonian Institution, Washington, D.C. 
20560. 


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CONTENTS 


Description of Eualus lineatus, new species, with a redescription of Heptacarpus herdmani 


(Walker) (Caridea: Hippolytidae) Mary K. Wicksten and Terrance H. Butler 
Hydroida from estuaries of South Carolina, USA: Families Sertulariidae and Plumulariidae 

Dale R. Calder 

Creedia alleni and Creedia partimsquamigera (Perciformes: Creediidae), two new marine fish 


species from Australia, with notes on other Australian creediids Joseph S. Nelson | 


Teleost fish remains (Osteoglossidae, Blochiidae, Scombridae, Triodontidae, Diodontidae) from 
the Lower Eocene Nanjemoy Formation of Maryland , 
Robert E. Weems and Stephen R. Horman 
Redescription of the Bigeye Shiner, Notropis boops (Pisces: Cyprinidae) 

Brooks M. Burr and Walter W. Dimmick 

Synonymy and distribution of Phyllomedusa boliviana Boulenger (Anura: Hylidae) 
David C. Cannatella 
Studies of the coastal marine fauna of southern Sinaloa, Mexico. IV. Report on the caridean 
crustaceans M. E. Hendrickx, M. K. Wicksten, and A. M. van der Heiden 
Three new species of Ochrotrichia (Metrichia) from Chiapas, Mexico (Trichoptera: Hy- 


droptilidae) Joaquin Bueno-Soria | 


Streptospinigera heteroseta, a new genus and species of Eusyllinae (Polychaeta: Syllidae) from 
the western shelf of Florida Jerry D. Kudenov 
The identity of Petrolisthes marginatus Stimpson, 1859, and the description of Petrolisthes 
dissumulatus, n. sp. (Crustacea: Decapoda: Porcellanidae) Robert H. Gore 
Geographic variation in Chlorospingus ophthalmicus in Colombia and Venezuela (Aves: 
Thraupidae) Storrs L. Olson 
Function of the teeth and vestibular organ in the Chaetognatha as indicated by scanning electron 
microscope and other observations Robert Bieri, Dolores Bonilla, and Fernando Arcos 
Systematics and distribution of shrews of the genus Crocidura (Mammalia: Insectivora) in 
Vietnam Lawrence R. Heaney and Robert M. Timm 
Parvulodesmus prolixogonus, a new genus and species of xystodesmid milliped from South 
Carolina (Polydesmida) Rowland M. Shelley 
Curidia debrogania, a new genus and species of amphipod (Crustacea: Ochlesidae) from the 
barrier reefs of Belize, Central America James Darwin Thomas 
Proechimys semispinosus (Rodentia: Echimyidae): Distribution, type locality, and taxonomic 
history Alfred L. Gardner 
The assignment of the Texas troglobitic water slater Caecidotea pilus to the genus Lirceolus, 
with an emended diagnosis of the genus (Crustacea: Isopoda: Asellidae) Julian J. Lewis 
Caecidotea fonticulus, the first troglobitic asellid from the Ouachita Mountains (Crustacea: 
Isopoda: Asellidae) Julian J. Lewis 
Psychronaetes hanseni, a new genus and species of elasipodan sea cucumber from the eastern 
central Pacific (Echinodermata: Holothuroidea) David L. Pawson 
Life diagram patterns in benthic polychaetes Kristian Fauchald 
Argyrotheca arguta, a new species of brachiopod from the Marshall Islands, western Pacific — 
Richard E. Grant 
Harbansus slatteryi, a new species of myodocopine ostracode from the Great Barrier Reef of 
Australia (Philomedidae) Louis S. Kornicker 


103 
110 
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134 
145 
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160 


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181 


See poe ee 


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“ i ocontings 
a of the 


BIOLOGICAL SOCIETY 
of 
WASHINGTON 


~ Volume 96 7 July 1983 Number 2 


THE BIOLOGICAL SOCIETY OF WASHINGTON 


1983-1984 
Officers 
President: David L. Pawson Secretary: Catherine J. Kerby 
Vice President: Donald R. Davis Treasurer: Leslie W. Knapp 
- Elected Council 
Darryl P. Domning C. William Hart 
Carl Ernst Robert P. Higgins 


David A. Nickle 


Custodian of Publications: Michael J. Sweeney 


PROCEEDINGS 
Editor: Brian Kensley 


Associate Editors 


Classical Languages: George C. Steyskal 3 Invertebrates: Thomas E. Bowman 
Plants: David B. Lellinger Vertebrates: Richard P. Vari 


Insects: Robert D. Gordon 


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Manuscripts, corrected proofs, editorial questions should be sent to the Editor, Biological 
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PROC. BIOL. SOC. WASH. 
96(2), 1983, pp. 189-201 


THREE NEW SPECIES OF COLOBOMATUS (COPEPODA: 
PHILICHTHYIDAE) PARASITIC IN THE MANDIBULAR 
CANALS OF HAEMULID FISHES 


Roger F. Cressey and Marilyn Schotte 


Abstract.—Three new species of Colobomatus are described from the mandib- 
ular canals of three genera of American haemulid fishes. These are the first species 
of Colobomatus reported from the Haemulidae and are the first described with 
abdominal processes in the female. Males of two of the new species are described. 
One of the species (C. quadrifarius) is restricted to the Eastern Pacific and the 
other two (C. caribbei and C. belizensis) to the western Atlantic. Data are pre- 
sented to suggest that an optimum host size exists for parasite infestation, accom- 
panied by diminishing infestations in smaller and larger size groups. 


During an examination of fishes in Belize for a study of host-parasite relation- 
ships, a new copepod belonging to the genus Colobomatus was discovered in the 
mandibular canals of its host fish, Haemulon sciurus (Shaw). Subsequent ex- 
amination of 475 tropical and subtropical haemulid fishes (three genera and 32 
species) yielded two additional new species from the same site on the hosts. 
Twenty of the 32 fish species examined were infested with one of the three new 
species. 

Since Hesse’s designation of the genus in 1873, 34 species of Colobomatus 
have been reported. They are members of the poecilostomatoid family Philichthy- 
idae and are known only from the mucous and sensory canal systems of marine 
fishes. The females are typically highly modified with a variety of appendages 
and processes thought to be useful in maintaining position in the lateral-line canals 
(Izawa 1974). The males are much less modified and are smaller. Males of only 
nine species have been described thus far. 

Sekerak (1970) published the first report of Colobomatus found in the “‘dentary 
canals’’ of host fish. Essafi and Raibaut (1980) described a closely related genus 
Colobomatoides from the pre-opercular cephalic system. All of the copepods in 
the present study were located in the mandibular canals of the hosts. 


Methods and Materials 


Fresh and alcohol-preserved fish were examined by removing the skin of the 
ventral portions of the lower jaws and exposing the mandibular canals. Figure 1 
shows the typical position of the parasite in one of the chambers of the canal. In 
total, 238 copepods were recovered, 181 @ and 1 6 of Colobomatus belizensis, 
47 2 and 2 6 of C. quadrifarius and 7 2 of C. caribbei. Numbers preceding host 
names in Materials Examined sections indicate the number of infested fish. Table 
2 indicates numbers of fish examined. The abbreviation USNM refers to Smith- 
sonian Institution and SOSC to Smithsonian Oceanographic Sorting Center spec- 
imens now in the general collections of the Division of Fishes, Smithsonian In- 
stitution. Drawings were prepared with the aid of a Wild M20 compound 
microscope with drawing tube. 


190 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


— a 


oe sy \ 
an oe | J ip N 
BS | | 
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v= AG oe 
Pg Pee ME 
{ { ie SS 
SS SS 
WA SS 
AS ln 
<i = 
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Fig. 1. Ventrolateral view of Haemulon mandibular canal showing position of Colobomatus in 
Situ. 


Colobomatus quadrifarius, new species 
Figs. 2C4E 


Material examined.—Holotype 2 (USNM 190517), allotype ¢ (USNM 190518), 
paratypes 15 2 (USNM 190519-22) from the mandibular canals of 17 Orthopristis 
reddingi Jordan and Richardson (USNM 43247, 167149, 167570 and 176154) from 
Baja California and Bay of Guaymas, Mexico. Additional material—4 ° from 3 
Anisotremus davidsoni (Steindachner) (USNM acc. no. 294075 and SOSC Ref. 
nos. 290 and 586) from Sonora, Mexico; 7 2 from 5 A. dovii (Gunter) (USNM 
144347-8 and USNM acc. no. 294075) from Colombia and Panama (both Pacific); 
6 2 from 3 A. interruptus (Gill) (USNM 181330) from Nayarit, Mexico; 3 2 and 
1 ¢ from 2 A. pacifici (Gunther) (USNM 114476, 220731) from El Salvador and 
Guatemala; 5 2 from 4 Haemulon flaviguttatum Gill (USNM 50426, 80548, 176149) 
from Panama and Baja California; 4 2° from 3 H. steindachneri (Jordan and 
Gilbert) (USNM 19632, 19879) from Colima and Cape St. Lucas, Mexico; 2 9 
from | Orthopristis chalceus (Gunther) (USNM 41389) from Galapagos. 

Description.—Female: Body form as in Fig. 2C. Length of body: 4.73—7.58 mm 
based on 3 specimens. Pre-oral cephalic process bifurcate. Body constricted be- 


Table 1.—Rate of infestation by Colobomatus belizensis in mandibular canals of 91 specimens of 
Haemulon sciurus related to size of fish. 


Host SL (mm) <80 80-109 110-139 140-169 170-199 200-229 >230 
Negative fish 13 7 2 0 2 Z 1 
One Colobomatus D) 5 D D 7 3 — 
Two Colobomatus 1 1 10 12 15 3 = 
Three Colobomatus 0 0 0 1 0 0 — 


% Infestation 19% 46% 86% 100% 92% 715% — 


~ 


VOLUME 96, NUMBER 2 191 


Fig. 2. A, Colobomatus caribbi, female, ventral; B, C. belizensis, female, ventral; C, C. quad- 
rifarius, female, ventral. 


192 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


tween thoracic segments 3 and 4. Thoracic segments 4 and 5 fused, bearing 2 
anteriorly directed and 2 posteriorly directed processes. In lateral aspect, pro- 
cesses usually extend from raised shoulders except in specimens from H. stein- 
dachneri. Sixth thoracic segment and genital segment (seventh according to Ka- 
bata 1979) each clearly separated. Four abdominal segments clearly separate, 
third bearing 2 conspicuous lateral processes. Caudal rami fused with last ab- 
dominal segment, about 8 times longer than wide. 

Male: Body form as in Figs. 3A and 3B. First antenna 6-segmented with setae 
as indicated in Fig. 3C. First 3 segments subequal in length, last three becoming 
progressively shorter. Second antenna (Fig. 3D) 5-segmented, first and second 
segments each with short spine, large recurved spines on each of last 3 segments 
and additional spine on terminal segment. 

Legs | and 2 biramous; rami somewhat overlapping. Leg | basipod with spine 
on outer distal corner; exopod (Fig. 4A) first segment with fringe on outer margin 
and stout pectinate spine on outer distal corner, last segment with 4 pectinate 
spines on outer margin and 3 inner plumose setae; endopod (Fig. 4B) first segment 
with inner plumose setae, last segment with 2 stout pectinate spines on outer 
distal margin and 4 inner plumose setae. Leg 2 exopod (Fig. 4C) first segment as 
in leg 1, last segment with 2 pectinate spines, distal fringed spine, and 2 inner 
plumose setae; endopod (Fig. 4D) as in leg | except outer margin of first segment 
with row of long hairs and only 3 plumose setae on inner margin of last segment. 
Leg 3 (Fig. 4E) represented by 5 setae, innermost 2 pectinate. 

Etymology.—The specific name (quadrifarius) is Latin meaning “‘four-fold”’ 
and refers to the four abdominal processes in the female. 

Remarks.—The female of C. quadrifarius is distinguished from its congeners 
by the presence of the long lateral processes on the third abdominal segment. 
None of the previously described species of Colobomatus has abdominal pro- 
cesses in addition to the caudal rami, although they are present in a closely related 
genus, Colobomatoides, described by Essafi and Raibaut (1980). 


Colobomatus belizensis, new species 
Figs. 2B, 4A—D, 6B-E, 7A—C 


Material examined.—Holotype 2 (USNM 190523) collected by the first author 
from mandibular canals of Haemulon sciurus (Shaw) 21 April 1981 at Carrie Bow 
Cay, Belize; allotype ¢ (USNM 190524) from the same host (USNM 167633) from 
Florida and 91 paratype 2 (USNM 190525—42) from the same host (USNM 15815, 
53293, 74404, 83798, 143731, 167610, 167633, 170872) from Florida (all locations 
in the Florida Keys, including Dry Tortugas except one from Pensacola); Belize 
(USNM 198808 and hosts examined at Carrie Bow Cay); Mexico (USNM 37089 
Cozumel, 192263 Yucatan); West Indies (USNM 34948, 38666, 38739); Cuba 
(USNM 192263). Additional material.—12 2 from 7 Haemulon aurolineatum Cu- 
vier (USNM acc. no. 261501, USNM 17716, 2 uncatalogued specimens) from 
South Carolina, Florida, and Brazil; 5 2 from 2 H. carbonarium Poey (USNM 
80586, USNM acc. no. 249542) from Dominica and Panama; 15 2 from 8 H. 
chrysarygyreum Giinther (USNM 6940, 35149-50, 35093, 41314) from Key West, 
Barbados, Abaco, and St. Lucia; | 2 from 1 H. macrostomum Gitnther (USNM 
80589) from Panama; 6 2 from 4 H. melanurum (Linnaeus) (USNM 3118, 10511, 


VOLUME 96, NUMBER 2 193 


Fig. 3. Colobomatus quadrifarius, male: A, Dorsal; B, Lateral; C, First antenna; D, Second 
antenna. 


194 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 4. Colobomatus quadrifarius, male: A, Leg 1; B, Leg 1, endopod; C, Leg 2; D, Leg 2, 
endopod; E, Leg 3. 


38479, 39808) from Bahamas, West Indies and Guyana; 2 2 from 1 A. parrai 
(Desmarest) (USNM 81097) from Toro Point, Panama; 20 2° from 12 AH. plumieri 
(Lacépéde) (USNM 37088, 82448, 142536, acc. no. 294075) from Cuba, Virgin 
Islands, Colombia (Caribbean) and Cozumel, Mexico; 16 @ from 10 H. stein- 
dachneri (Jordan and Gilbert) (USNM 10265, acc. no. 278057, Oregon stations 
5654, 10536, 17628-9) from Caribbean coast of South America (Colombia to Bra- 
zil); 9 2 from 9 Orthopristis chrysopterus (Linnaeus) (USNM 35205, 163701) 


VOLUME 96, NUMBER 2 195 


Fig. 5. Colobomatus belizensis, female, SEM photos: A, Oral area; B, Labrum; C, Tip of second 
maxilla; D, Second maxilla in situ within oral cone (lower fringe-labium?). 


North Carolina and Louisiana; 3 @ from 2 O. ruber (Cuvier and Valenciennes) 
(USNM 123123, 278057) from Venezuela and Guyana. 

Description.—Female: Body form as in Fig. 2B. Length of body: 4.13-9.38 
mm, based on 3 specimens. Single, simple cephalic lobe, acute at tip, extending 
above buccal cone. Degree of segmentation at base of third segment (“‘neck’’) 
variable. Four long, slender processes, 2 anterior and 2 posterior, simple and 
usually rounded at tip. Genital segment (seventh thoracic segment) with small 
lateral lobes. Egg sacs often as long as abdomen, easily broken when copepod is 
handled. Four abdominal segments more or less definite; penultimate segment 
with 2 lateral processes. Caudal rami well-developed and similar to those pro- 
cesses described above. 

First antenna very small, placed dorsolaterally near insertion of buccal cone. 
Oral area typical of genus (see SEM photos, Figs. 5A—D). Labrum modified as 
forked, posteriorly directed process between mandibles. Mandibles with small 
ventral process; first maxilla bifid. Second maxilla considerably larger, bifid, ter- 
minal segments with numerous large spinose processes. Maxillipeds absent. 


196 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 6. Colobomatus quadrifarius, female: A, ventral; C. belizensis: B—E, Ventral. 


VOLUME 96, NUMBER 2 


Fig. 7. Colobomatus belizensis, 


male: A, First antenna; B, Leg 1; C, Leg 2. 


197 


198 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Male: Total length 1.94 mm. Width of cephalon 0.36 mm. General body form 
typical of genus with wide cephalon, body tapering gradually to end of abdomen 
bearing elongate, caudal rami. Third thoracic segment with 2 dorsolateral processes 
directed posteriorly with tips curving away from body. 

First antenna (Fig. 7A) 6-segmented with one aesthete on each of last 3 seg- 
ments; first 4 subequal in length. Second antenna similar to that of C. quadrifarius 
(Fig. 3B). 

Legs 1 and 2 biramous (Figs. 7B and 7C) with patch of spinules on basipods. 
The 2-segmented exopod of leg 1 with one stout spine on first segment and 4 on 
the second, all with numerous spinules along both margins. Second segment 
bearing 3 long plumose setae; setae also found on interior margins of both seg- 
ments. 

Short first segment of endopod unarmed; second segment, 2.5 times as long, 
bearing 2 strongly sclerotized spines, innermost twice as long as outermost, each 
bearing spinules on margins. 

Leg 2 with one less spine on exopod than leg |. Endopod with 2 setae and 3 
spines. Leg 3 reduced to 5 setae, similar to that of C. quadrifarius (Fig. 4E). 
Fifth thoracic segment bearing 2 lateral setae. Each caudal ramus bearing one 
lateral seta and 4 terminal setae. 

Etymology.—The specific name, belizensis, is derived from the type-locality. 

Remarks.—The female of C. belizensis is similar in body form to C. quadri- 
farius and like it and the other new species, C. caribbei, can be differentiated 
from others in the genus by the presence of two long lateral processes extending 
from the penultimate abdominal segment. The present species has a single, sim- 
ple, cephalic projection anterior to the oral cone in contrast to the bifurcate 
process of C. quadrifarius. The thoracic processes appear to be more dorsally 
placed as well as raised in C. quadrifarius. 


Colobomatus caribbei, new species 
Fig. 2A 


Material examined.—Holotype 2 (USNM 190543) from mandibular canal of 
Anisotremus surinamensis (Bloch) (USNM 123121) from Venezuela, paratypes 6 
2 (USNM 19054446) from the same host (USNM 30878, 80533, 123121) from 
Florida, Panama, and Venezuela. 

Description.—Female: Body form as in Fig. 2A. Body length: 4.95-12.23 mm 
based on 5 specimens. Anterior cephalic process bifid. Head and first 3 thoracic 
segments fused. Thoracic segments 4 and 5 separated from anterior segments by 
constriction and bearing 2 anteriorly directed processes and 2 laterally directed 
processes; processes bulbous in basal half and attenuated in distal half. In dorsal 
aspect, the processes appear to be borne on raised shoulders. Thoracic segments 
6 and 7 (genital segment) distinct. First 2 abdominal segments distinct. Abdominal 
segments 3 and 4 fused; segment 3 with 2 lateral processes. Caudal rami atten- 
uated in distal half. 

Male: Unknown. 

Etymology.—The specific name, caribbei, is derived from the type-locality in 
the Caribbean Sea. 

Remarks.—As in the two previous species, this species is characterized by the 


VOLUME 96, NUMBER 2 199 


Table 2.—Synopsis of hosts examined and Colobomatus infestations of each (A = Atlantic, P = 
Pacific). 


No. of fish No. of fish Colobomatus 

Fish species examined infested % infested spp. 
Anisotremus caesius (P) 4 0 0 neg 
A. davidsoni (P) 4 3 75 quadrifarius 
A. dovii (P) 8 5 60 quadrifarius 
A. interruptus (P) 10 3 33 quadrifarius 
A. pacifici (P) 12 2 17 quadrifarius 
A. surinamensis (A) 9 4 44 caribbei 
A. virginicus (A) 15 0 0 neg 
Haemulon album (A) 11 0 0 neg 
H.. aurolineatum (A) 14 7 50 belizensis 
H.. bonariense (A) 17 0 0 neg 
H. boschmae (A) 26 0 0 neg 
H. carbonarium (A) 13 2 15 belizensis 
H. chrysargyreum (A) 9 8 89 belizensis 
H. flaviguttatum (P) 13 4 3] quadrifarius 
H. flavolineatum (A) 15 0 0 neg 
H. macrostomum (A) 13 1 8 belizensis 
H. maculicauda (P) 16 0 0 neg 
H. melanurum (A) 10 4 40 belizensis 
H. parrai (A) 13 1 8 belizensis 
H. plumieri (A) 13 12 92 belizensis 
H. sciurus (A) 91 64 70 belizensis 
H. scudderi (P) 14 0 0 neg 
H. sexfasciatum (P) 10 0 0 neg 
H. steindachneri (A) 21 10 45 belizensis 
H. steindachneri (P) 16 3 19 quadrifarius 
H. striatum (A) 16 0 0 neg 
Orthopristis brevipinnis (P) 3 0 0 neg 
O. chalceus (P) 12 1 8 quadrifarius 
O. chrysopterus (A) 6 5 83 belizensis 
O. forbesi (P) 2 0 0 neg 
O. reddingi (P) 30 17 56 quadrifarius 
O. ruber (A) 9 2 22 belizensis 


presence of lateral processes on the penultimate segment. The bifid cephalic lobe 
distinguishes the present species from C. belizensis, and the bulbous nature of 
the appendages as well as the fused abdominal segments separate it from C. 
quadrifarius and C. belizensis. 


Discussion 


The three new species of Colobomatus described here appear to be closely 
related on the basis of the presence of the two lateral processes on the third 
abdominal segment of each. Essafi and Raibaut (1980) described a new genus 
Colobomatoides based in part on the presence of abdominal processes in the 
female. Although the three species described here possess abdominal processes, 
we feel that sufficient differences exist between Colobomatoides and these new 
species to warrant including the three new ones in the genus Colobomatus. These 


200 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Colobomatus belizensis 
eC’. caribber 


C. quadrifarius 


120. NS ike) 105 lfele} 95 


Fig. 8. Distribution of three new species of Colobomatus. 


new species, however, are the first Colobomatus species with this character to 
be described. 

The generic diagnosis of Colobomatus should be amended to state that the 
female may or may not possess abdominal processes and the key to philichthyid 
genera provided by Essafi and Raibaut (1980) modified. 


Effect of Host Size 


Table | suggests that an optimum size host is preferred by the parasite. All 15 
specimens of Haemulon sciurus between 140-169 mm SL were infested. The rate 
of infestation diminishes in smaller and larger size groups. 


Host Specificity and Biogeography 


The present work includes species of hosts from three haemulid genera present 
in the tropical and semitropical western Atlantic and eastern Pacific (see Table 
2). Figure 8 shows the distribution of each of the species. Colobomatus quadri- 
farius infests species of three genera (Haemulon, Orthopristis, and Anisotremus) 
in the eastern Pacific with little host specificity indicated beyond a possible re- 
striction to haemulid fishes. The two Atlantic Colobomatus species, however, do 
show some degree of host specificity as C. caribbei is known thus far only from 
Anisotremus, and C. belizensis from Haemulon and Orthopristis. 


Acknowledgments 


The work in Belize was facilitated by Dr. Klaus Ruetzler (Smithsonian Insti- 
tution), director of the Investigations of Marine Shallow Water Ecosystems proj- 


VOLUME 96, NUMBER 2 201 


ect at Carrie Bow Cay, Belize. This paper is contribution number 119 of the 
Investigations of Shallow Water Ecosystems Project, supported in part by the 
Exxon Corporation. 


Literature Cited 


Essafi, K., and A. Raibaut. 1980. Colobomatoides splendidus n.g., n.sp., (Copepoda, Philichthyi- 
dae) parasite de poissons téléostéens du genre Sparus (Sparidae) des cétes de Tunisie.—Extrait 
des Archives de |’Institut Pasteur de Tunis 57(4):355-361. 

Izawa, K. 1974. On three new species of Colobomatus (Cyclopoida: Philichthyidae) parasitic on 
Japanese fishes.—Publications of the Seto Marine Laboratory 21:335—343. 

Kabata, Z. 1979. Parasitic copepods of British fishes. 468 pp. London: The Ray Society. 

Sekerak, A. 1970. Parasitic copepods of Sebastes alutus including Chondracanthus triventricosus 
sp. nov. and Colobomatus kyphosus sp. nov.—Journal Fisheries Board of Canada 27 (11): 
1943-1959. 


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


PROC. BIOL. SOC. WASH. 
96(2), 1983, pp. 202-224 


NOTES ON THE BIOLOGY OF SOME 
SEAGRASS-DWELLING CRUSTACEANS 
(STOMATOPODA AND DECAPODA) 
FROM CARIBBEAN PANAMA 


Loren D. Coen and Kenneth L. Heck, Jr. 


Abstract.—A year-long sampling program in seagrass meadows along the Ca- 
ribbean coast of Panama yielded two stomatopod and 58 decapod species in 45 
genera and 22 families, a number of which was previously unreported from Pan- 
amanian waters. Though not complete, this annotated list provides information 
on size, seasonality, reproductive state, and presence of parasites. High species 
affinities with recent collections from Bermuda and the Carolinas exist. Specifi- 
cally, 59% of the species treated here are reported from the Carolinas and 64% 
from Bermuda; however, no regional endemics were found. Range extensions 
are made for several species. Recent literature and taxonomy are reviewed for 
most species discussed. 


Abele, in a symposium volume on the Panamic biota in 1972, pointed out that 
almost no information was available at that time on the crustaceans of Panama. 
In recent years there have been publications on the brachyuran fauna of the 
Bahamas (Garth 1978) and the north coast of Colombia (Lemaitre 1981), and on 
the decapod fauna of Bermuda (Markham and McDermott 1981). Yet, with the 
exception of Gore and Abele’s (1976) publication on the porcellanid crabs of 
Panama, and Gore’s (1982) recent study on the Porcellanidae of Central America, 
our knowledge of the Panamanian crustacean fauna, especially on the Caribbean 
coast, remains nearly as limited as when Abele made his comments ten years 
ago. 

Here we provide new information on a number of poorly known decapod and 
stomatopod crustaceans found primarily in seagrass habitats along the Caribbean 
coast of Panama. Included are data on size, reproductive condition, habitat, and 
other observations on the natural history of the 60 species treated here. This is 
clearly a small percentage of the total decapod and stomatopod fauna of the 
Caribbean coast of Panama and our list is in no way comprehensive. However, 
we believe that it does provide an indication of the species commonly found in 
subtidal seagrass habitats along the Caribbean Panamanian coast. Other infor- 
mation on the species treated here has been previously published by Heck (1977, 
1979) and Heck and Wetstone (1977). 


Study Sites 


Sites consisted of vegetated subtidal meadows within 30 m of the shore. Study 
areas were located 20 km to the northeast of the Caribbean end of the Panama 
Canal (Fig. 1). Detailed descriptions of our stations 1-4 are provided by Heck 
(1977) and Weinstein and Heck (1979). Station 1 was seaward of a thick mangrove 
shoreline (Rhizophora mangle (L.)) and contained thick growths of Thalassia 


VOLUME 96, NUMBER 2 203 


BR | ; 
3 fie ee: CARIBBEAN 
ne SEA 


GALETA 
LAB 


Stud 
Area 


Fig. 1. Location of collection sites along the Caribbean coast of Panama. 


testudinum (Konig) and Halimeda opuntia (L.) with lesser amounts of Syringo- 
dium filiforme (Kutzing), Udotea flabellum (F.) (Ellis and Solander) and Peni- 
cillus capitatus (Lamarck). Sediments were mostly fine muds, and the site was 
protected from wave shock by a small patch reef. Station 2 was exposed to the 
most severe wave shock of our 5 study areas. Dominant vegetation included T. 
testudinum and S. filiforme. Sediments were calcareous sand with some larger 
coral and Halimeda fragments. Station 3 received the least wave shock among 
our 5 sites. Thalassia testudinum was the dominant plant, although H. opuntia 
and H. incrassata (Ellis) were also present. Sediments were coarse with many 
coral and Halimeda fragments. Station 4 was surrounded by reefs and located in 
the lee of two small islands. Thalassia testudinum was the dominant plant al- 
though S. filiforme was also present. Sediments were coarse with many coral 
fragments. Station 5 was located in a small cove lined with red mangroves (R. 
mangle). Vegetation was sparse, with some 7. testudinum and S. filiforme pres- 
ent. 

Salinities ranged from nearly 0% during heavy rains to nearly 38% during the 
dry season, although values were most often similar to open sea salinities (~38%c). 
Water temperatures ranged from about 27-31°C (see Glynn 1972, for additional 
information on the physical regime). Although the stations were subtidal, water 
depth rarely exceeded 2 m or became shallower than | m during any one tidal 
cycle. 


Methods 


Samples were collected by a 4.87 m otter trawl, with 19 mm stretch mesh wings 
and a 6.3 mm liner. Monthly samples (station 1-4) were taken during the period 
of July 1974-May 1975, according to the schedule described by Heck (1977). 
Station 5 was only sampled during July, August, and September. Samples were 


204 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


taken diurnally, except for one set of night samples in May 1975 at stations 1, 2 
and 3 (an average of 4 trawls per station). All samples were preserved in 10% 
formalin and later transferred to 70% isopropanol before identifications were made. 
All available specimens were identified, sex determined and measured to gather 
information on size at sexual maturity. By our criteria male shrimps were mature 
if the appendix masculina on the second pleopod was apparent (with a few ex- 
ceptions, e.g., Synalpheus in which the appendix masculina is absent). Female 
shrimps were only separated into ovigerous and non-ovigerous categories. Ma- 
turity of anomuran and brachyuran crabs follows Abele and Blum’s (1977) cri- 
teria. Male crabs were considered mature with the development of gonopods; 
females if the abdomen was covered by the sternum. 

Most measurements are given for carapace length (cl), measured by dial caliper 
to the nearest 0.1 mm from posterior margin of the carapace to the postorbital 
margin. For crabs, carapace width (cw) includes the greatest width measured 
including spines. For hermit crabs, shield length (sl) was measured as in Proven- 
zano (1959). Descriptions were not considered necessary but recent faunal sum- 
maries and classic monographs where these may be found are summarized (Table 
1) as an aid to future works in the zoogeographical areas. The classification and 
nomenclature scheme used here has been adopted from The Biology of Crustacea 
(1982; L. G. Abele, pers. comm.) and Abele and Felgenhauer (1982). 

Part of the collection was lost or badly damaged in transport from Panama, 
and resulting deficiencies are noted where appropriate in the text. Thus, many of 
the preliminary identifications (Heck 1977) done by authorities in the field are 
retained, though the specimens could not be measured or sexed. 


ANNOTATED SPECIES LIST 


SUPERORDER HOPLOCARIDEA 
ORDER STOMATOPODA 
Family Gonodactylidae 


Gonodactylus lacunatus Mannins, 1966 


Material.—Station 5; Sept; 1 male. 

Measurements.—cl 5.4 mm. 

Distribution.—From the western Atlantic, the Caribbean Sea, Yucatan, Nica- 
ragua and Colombia; sublittoral to 50 m, though usually in shallower waters. 


Family Pseudosquillidae 
Pseudosquilla ciliata (Fabricius, 1787) 


Material.—Stations 1, 3 and 4; Aug, Sept, Oct, Feb and Apr; 7 individuals. 

Measurements.—Males, 4.3 to 7.9 mm cl; females, 5.0 to 7.3; juveniles, 3.4 to 
355). 

Habitat.—Shelly-sand, coralline algae, coral, though most often seagrasses. 

Distribution.—From tropical oceans in the Indo-West Pacific, the western At- 
lantic, Bermuda, the Bahamas, Florida and western Africa; to 110 m, though 
primarily in shallower waters. 


VOLUME 96, NUMBER 2 


205 


Table 1.—Brief summary of the carcinological literature (Decapoda) relating to the faunal area.* 


Author(s) 


FAUNAL SUMMARIES 


Abele 1970 
Abele 1972a,b 
Felder 1973 


Garth 1978 
Heck 1977 
Lamaitre 1981 


Markham and McDermott 1981 


Park 1969 
Powers 1977 
Rouse 1970 
Wass 1955 
Williams 1965 


# of 
species 
treated? 


154 
485 
143 


53 
65 
96 
276 
15 
352 
103 
113 
220 


Taxonomic group discussed 


Decapoda of northwestern Gulf of Mexico 


Pacific and Caribbean Decapods of Panama 


Crabs and lobsters of the northwest Gulf 
of Mexico 

Brachyura of the Bahamas 

Decapoda, Caribbean Panama 

Shallow-water Brachyura of Colombia 

Decapoda of Bermuda 

Portunidae of Biscayne Bay 

Crabs of the Gulf of Mexico 

Littoral Crustacea of southwest Florida 

Decapoda of northwest Florida 

Decapoda of the Carolinas 


TAXONOMIC WORKS (including monographs) 


Camp 1973 
Chace 1972 


Coutiere 1909 
Gore and Abele 1976 


Gore and Scotto 1979 
Haig 1960 

Holthuis 1958 
Holthuis 1959 
Lemaitre et al. 1982 
Lyons 1970 


Manning 1961 
Manning 1969 


Manning and Chace 1971 


Perez Farfante 1969 
Perez Farfante 1971 
Provenzano 1959 
Rathbun 1918 
Rathbun 1925 
Rathbun 1930 
Rathbun 1937 
Williams 1974b 


28 


218 


23 
238 
273 
349 
127 

14 


Stomatopoda of Florida (Hourglass 
Cruises) 

Shrimps of the Smithsonian-Bredin Expe- 
dition (Caribbean and West Indies) 

American Synalpheus species 

Pacific Panama and adjacent 
Caribbean, Procellanidae 

Western Atlantic Parthenopidae 

Eastern Pacific Porcellanidae 

Calappidae of the West Indies 

Decapoda of Suriname (Dutch Guiana) 

Provenzanoi group of pagurid crabs 

Scyllaridae of Florida (Hourglass 
Cruises) 

Genus Leander 

Stomatopoda of the western Atlantic 

Processidae of the northwestern Atlantic 

Genus Penaeus 

Genus Metapenaeopsis 

Shallow-water hermit crabs of Florida 

Grapsoid crabs of America 

Spider crabs of America 

Cancroid crabs of America 

Oxystomatous crabs and allied groups 

Genus Callinectes 


Comments 


> 


ey 


ppp Pp P >P HH p 
nn A 


& 
= 


ie 
ap kar 


ppp ppp 
FAA A 


ppeppeepepppep Pp p 
RRA RRRRAARAKR 


ey ey s . . 


ey 


* NOTE: This is not meant to be exhaustive, but rather a guide to the most accessible literature; 
earlier references may be omitted if they are covered within the scope of another, more recent 
treatment. Totals may be approximate to account for supergenera, subspecies, etc. 

+ Total number of species addressed (includes subspecies and those reviewed from adjacent waters). 

+ Key to contents of papers: a, annotated list; f, faunal list without additional information; k, keys 


included (or new descriptions). 


206 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


ORDER DECAPODA 
SUBORDER DENDROBRANCHIATA 
SUPERFAMILY PENAEOIDEA 
Family Penaeidae 


Metapenaeopsis martinella Perez Farfante, 1971 


Material.—Stations 1, 2, 3 and 5; Sept and May; 10 individuals. 

Measurements.—One juvenile male, 10 mm cl; mature females, 9.8 to 12.6. 

Habitat.—Chace (1972) reported specimens from coral reefs, shelly bottoms 
and calcareous algae. 

Remarks.—This species was identified in Heck (1977) as Trachypenaeus res- 
trictus but reexamination allowed the identification to be corrected. Most indi- 
viduals from night samples, stations 2 and 3. The species known from Cuba, 
Brazil, and the western Caribbean; 4 to 137 m. Genus reviewed by Perez Farfante 
(1971). 


Penaeus (Melicertus) duorarum notialis Perez Farfante, 1967 


Material.—Stations 1—5; present in all sampled months; 300 individuals. 

Measurements.—Males, 3.9 to 6.3 mm cl; females, 4.5 to 20.6. 

Habitat.—On muds, sand and sand within rocky patches. 

Remarks.—Specimens taken primarily in day trawls (cf. Greening and Living- 
ston 1982); most were immature. Over 80% from station 1. Subspecies ranges 
from Cuba to the Virgin Islands, Mexico, the Caribbean Sea to Brazil and the 
western coast of Africa; to 732 m, though generally to 65 m. Genus reviewed by 
Perez Farfante (1969). 


Family Sicyonidae 
Sicyonia laevigata Stimpson, 1871 


Material.—Stations 1, 2 and 3; May only; 16 individuals. 

Measurements.—Males (two), 7.4 mm cl; mature females, 7.2 to 11.1. 

Habitat.—Common in areas with abundant shell cover and among rocks; also 
in Thalassia. 

Remarks .—All but two (87%) collected in night samples. Williams (1965) noted 
sexually mature individuals as small as 18 mm total length. Greening and Living- 
ston (1982) reported S. /aevisata in their nighttime seagrass samples. Species 
ranges from North Carolina to northwestern Florida, the West Indies, Colombia 
and the Pacific coast of Panama; to 90 m. 


SUBORDER PLEOCYMATA 
INFRAORDER STENOPODIDEA 
Family Stenopodidae 


Stenopus hispidus (Oliver, 1811) 


Material.—Station 3; Sept and May (night); 2 individuals. 
Measurements.—Both males, 6.2 and 7.6 mm cl. 
Habitat.—Grass flats and areas with coral rubble (Chace 1972). 


VOLUME 96, NUMBER 2 207 


Distribution.—Widely ranging from Bermuda, central eastern Florida and south 
Florida to the West Indian region, Surinam, the Red Sea, Japan, southeast Africa, 
and Hawaii; to 210 m. 


INFRAORDER CARIDEA 
SUPERFAMILY PALAEMONIDEA 
Family Palaemonidae 


Leander tenuicornis (Say, 1818) 


Material.—Stations 1-3; April and May; 24 individuals. 

Measurements.—Males, 4.9 to 5.1 mm cl; ovigerous females, 6.3 to 8.4. 

Habitat.—Most often associated with vegetation, Sargassum, Thalassia and 
mangroves (Ledoyer 1969; Chace 1972). 

Remarks.—Most individuals from station 1. It is distributed in shallow or pe- 
lagic waters of all tropical and subtropical seas except those in the extreme east- 
ern Pacific. Only 2 species known in American waters. See Manning (1961) for a 
comparison of this species with L. paulensis (Ortmann). 


Periclimenes americanus (Kingsley, 1878) 


Material.—Majority at station 1; collections scattered throughout sampling pe- 
riod; 19 individuals. 

Measurements.—Males, 2.3 to 3.5 mm cl; ovigerous females, 2.7 to 3.3. 

Habitat.—Associated with coral reefs, on mud and sand flats, submerged struc- 
tures and especially in seagrass and mangrove habitats (Chace 1972). 

Distribution.—Species ranges from Bermuda, North Carolina to northwest 
Florida, Yucatan Peninsula, and West Indies; to 73 m. 


Family Gnathophyllidae 
Gnathophylloides mineri Schmitt, 1933 


Material.—One ovigerous female; data lost. 

Measurements.—2.2 mm cl. 

Habitat.—Among sea urchin spines, rocks, and coral rubble (Chace 1972). 

Distribution.—From Bermuda, southeastern Florida, Yucatan Peninsula and 
the Caribbean Sea; littoral and sublittoral. 


SUPERFAMILY ALPHEOIDEA 
Family Processidae 


Ambidexter symmetricus Manning and Chace, 1971 


Material.—Stations 1, 2 and 3; May; 7 individuals. 

Measurements.—Males, 3.4 to 4.3 mm cl; ovigerous females, 3.4 to 5.2. 

Habitat.—Common on vegetated bottoms, especially seagrasses. 

Remarks.—One individual (5.8 mm cl) was infested with a bopyrid isopod 
(branchial; Table 2). Processid shrimp are a common seagrass-inhabiting group, 
being taken especially at night (Ledoyer 1969; Manning and Chace 1971; Greening 
and Livingston 1982; personal observations). This species is noted from the west- 


208 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


ern Atlantic, including eastern and western Florida (Gulf of Mexico), Puerto Rico 
and Trinidad; to 6 m. This is the first report of A. symmetricus from Panama. 


Processa fimbriata Manning and Chace, 1971 


Material.—Most from stations 1 and 2; Apr and May; 55 individuals. 

Measurements.—Males, 2.9 to 4.2 mm cl; ovigerous females, 5.4 to 5.6. 

Habitat.—Areas of broken shell, coral rubble, sponges and Thalassia. 

Remarks.—The majority of the individuals collected at night (93%). Reported 
from North Carolina to Brazil, including south Florida, the Bahamas and Puerto 
Rico; sublittoral to 37 m. This constitutes a new record for Panama. 


Family Alpheidae 
Alpheus armillatus H. Milne Edwards, 1837 


Material.—Stations 1, 2 and 3; May only; 16 individuals. 

Measurements.—Males, 5.3 to 9.2 mm cl; one non-ovigerous female, 11.7; 
ovigerous females, 5.3 to 11.7. 

Habitat.—In oyster bars, coral rubble, rocks and Thalassia beds (Chace 1972). 

Distribution.—Bermuda, North Carolina to the Gulf of Mexico and Brazil; 
sublittoral. 


Alpheus floridanus Kingsley, 1878 


Material.—Stations | and 2; Oct, Dec and Jan; 8 individuals. 

Measurements.—Males, 4.3 to 8.8 mm cl; ovigerous females, 6.4 to 19.8. 

Habitat.—Soft-sediments. 

Distribution.—From Bermuda, the southeastern United States to the Gulf of 
Mexico, Brazil, Guyana and the Congo (eastern Atlantic); to 37 m. 


Alpheus formosus Gibbes, 1850 


Material.—Stations 1 and 3; Aug and Mar; 2 females. 

Measurements.—3.6 and 5.5 mm cl. 

Habitat.—On sand and mud flats, rocks, oyster bars and coral rubble. 

Distribution.—From Bermuda, North Carolina and throughout the West Indies 
to Brazil; to 42 m. 


Alpheus normanni Kingsley, 1878 


Material.—Stations | and 5; Oct and May; 4 individuals. 

Measurements.—Males, 4.1 to 5.4 mm cl; one ovigerous female, 6.3. 

Habitat.—In Thalassia, sponges and coral rubble (especially Porites); also 
among ascidians (Abele 1970). 

Remarks.—All 3 males were found associated with a sponge (Station 5). Wass 
(1955) found this shrimp to be common in the northeast Gulf of Mexico, and 
Greening and Livingston (1982) collected this species mostly at night. Found from 
Bermuda, Virginia to the Gulf of Mexico, throughout the West Indies to Tobago 
and in the Pacific in the Gulf of California; littoral to 73 m. 


VOLUME 96, NUMBER 2 209 


Synalpheus fritzmuelleri Coutiére, 1909 


Material.—Stations 1 and 2; Aug, Feb and May; 6 individuals. 

Measurements.—Ovigerous females (two), 3.2 and 5.0 mm cl; remaining spec- 
imens, 2.9 to 3.8. 

Habitat.—Commonly collected in sponges (Abele 1970) and in grass flats, as 
well as among mangrove roots (Chace 1972). 

Remarks.—Both ovigerous females collected from sponges. S. fritzmuelleri 
recorded from Bermuda, North Carolina to the Gulf of Mexico and the West 
Indies to Brazil; in the eastern Pacific from Baja, California; littoral to SO m. 


Synalpheus goodei Coutiere, 1909 


Material.—Station 2; Oct; 19 individuals. 

Measurements.—Ovigerous females (13), 4.6 to 6.1 mm cl. 

Habitats.—Associated with rocky intertidal areas encrusted with corals; Gore 
(pers. comm.) found it living on or near a large keratose sponge. 

Remarks.—Six of the 19 specimens lost in transit. Species known from Ber- 
muda, central east Florida Gore (pers. comm.), the Gulf of Mexico and Curag¢ao; 
to 60 m. 


Synalpheus pandionis Coutiere, 1909 


Material.—Stations 1 and 2; Aug and Feb; 3 individuals. 

Measurements.—Ovigerous females (three), 4.9 to 5.5 mm cl. 

Habitat.—Collected from Thalassia with clumps of Porites rubble and algae 
(Chace 1972). 

Distribution.—This rare species reported from the Gulf of Mexico, Barbados 
and Cura¢ao; to 60 m. 


Synalpheus townsendi Coutiére, 1909 


Material.—Station 2; Aug and Feb; 4 individuals. 

Measurements.—One ovigerous female, 4.0 mm cl; the remaining specimens, 
1.9 to 3.9. 

Habitat.—Common in sponges (Abele 1970) and Thalassia beds with clumps 
of Porites (Chace 1972). Gore (1981) recorded it from a deep reef in the Florida 
Keys. 

Remarks.—Chace (pers. comm.) suggests that references to associations with 
Pocillopora are in error, referring probably to growth forms of Porites. Species 
occurs from Bermuda and North Carolina, southwest through the Gulf of Mexico 
and the West Indies to Brazil; to 102 m. 


Family Hippolytidae 


Hippolyte zostericola (Smith, 1873) 


Material.—Stations 2 and 3; Sept and May; 4 individuals. 

Measurements.—Four ovigerous females, 1.7 to 2.6 mm cl. 

Habitat.—Usually common in submerged vegetation (Chace 1972; Greening 
and Livingston 1982). 


210 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Remarks.—From Bermuda and Massachusetts to the Yucatan Peninsula, Trin- 
idad and Curacao; in shallow waters. See Williams (1965) and Chace (1972) for 
a discussion of its taxonomic difficulties. 


Latreutes fucorum (Fabricius, 1798) 


Material.—Stations 1 and 2; Apr and May; 15 individuals. 

Measurements.—Ovigerous females, 2.3 to 3.5 mm cl; a single non-ovigerous 
female, 2.2. 

Habitat.—Most often associated with floating and submerged vegetation (Wass 
1955; Abele 1970; Chace 1972). 

Distribution.—Ranges from Newfoundland to the Gulf of Mexico, Bermuda 
and Puerto Rico; pelagic (in vegetation) and sublittoral. 


Thor manningi Chace, 1972 


Material.—Stations 1, 2 and 5; Aug and May; 12 individuals. 

Measurements.—Single male, 2.3 mm cl; ovigerous females, 1.9 to 3.7; females 
(non-ovigerous), 1.8 to 2.6. 

Habitat.—Common in grass flats. 

Remarks.—Most specimens were taken in night samples. Recorded from Ber- 
muda, North Carolina to Tobago, Curacao and in the eastern Pacific; to 4 m. 
Chace (1972) suggested that this hippolytid may be a sequential (protandric) her- 
maphrodite. 


Tozeuma carolinense Kinglsey, 1878 


Material.—Stations 1-5; all months sampled; 1068 individuals. 

Measurements.—Males (mature), 3.5 to 6.7 mm cl; ovigerous females, 5.1 to 
10.8; non-ovigerous females, 5.2 to 9.4. Several males less than 3.5 mm cl were 
collected, each with a developing appendix masculina. 

Habitat.—Common in vegetation, with color being highly variable (Abele 1970; 
Greening and Livingston 1982; pers. obs.). 

Remarks.—T. carolinense dominated with decapoda fauna (ranked by abund- 
ance) at all stations. Ovigerous females were found throughout the year, with 
ovigery in a given collection always greater than 65%. Previous studies on this 
species have been primarily concerned with its larval development (Bryce 1961; 
Ewald 1969), although Voss (1956) briefly discussed its natural history. Found 
from Bermuda and Massachusetts to Panama; littoral to 75 m. 


Trachycaris restrictus (A. Milne Edwards, 1878) 


Material.—Stations 2, 3 and 5; Aug, Mar, Apr and May; 13 individuals. 

Measurements.—Ovigerous females (13), 5.0 to 6.8 mm cl. 

Remarks.—Genus with one species. This rare and curious looking hippolytid 
has been recorded from Bermuda to Brazil and in the eastern Atlantic from 
Canary Islands to Saint Helena Island (Holthuis 1949); to 100 m. This is probably 
the first record of this species from Panama. 


VOLUME 96, NUMBER 2 211 


INFRAORDER PALINURA 
SUPERFAMILY PALINUROIDEA 
Family Scyllaridae 


Scyllarus Fabricius, 1775 


Remarks.—Two individuals were collected; however, both were lost in transit. 
See Williams (1965) and Lyons (1970) for a review of this group. 


Family Palinuridae 
Palinurus argus (Latreille, 1804) 


Material.—Stations 1, 2 and 5; Aug, Oct, Mar, Apr and May; 27 juvenile 
individuals. 

Measurements.—Specimens ranged from 11.2 to 37.4 mm cl. 

Habitat.—Spiny lobsters commonly found around rocky and coral reef areas; 
generally in areas offering concealment (Khandler 1964; Davis 1971, 1981; Berrill 
1975). 

Remarks.—Juveniles were taken primarily at Station 1, possibly associated 
with the nearby red mangroves, where dense prop roots offer areas of conceal- 
ment (Heck 1977). P. arsus has been recorded from Bermuda, North Carolina to 
the Gulf of Mexico, the West Indies and Brazil; to 100 m. Williams (1965) gives 
a general overview of this species’ ecology. 


INFRAORDER ANOMURA 
SUPERFAMILY COENOBITOIDEA 


(Note: hermit crabs were tentatively identified in Panama by one of us (KLH); 
however, most were damaged during extraction. Though intact specimens were 
originally verified, measurements and sexing for most specimens were impossi- 
ble). 


Family Diogenidae 
Calcinus tibicen (Herbst, 1791) 


Material.—Stations 2, 3 and 5; Sept and May night; 3 individuals. 

Measurements.—One specimen measured, 4.4 mm sl. 

Habitat.—Common in rocky shores. 

Remarks.—Found in Bermuda, the West Indies, Florida and Brazil; intertidal 
to 32 m. See Provenzano (1959) for further information. 


Clibanarius antillensis Stimpson, 1862 


Material.—Stations 1—5; all sampled months; 91 individuals. 

Measurements.—Measurable specimens from 1.6 to 4.7 mm sl. 

Remarks.—Ovigerous females were present in August and September. Over 
68% of the specimens from station 2. Known from Bermuda, south Florida, the 
West Indies, Curacao and Brazil; in shallow waters. See Provanzano (1959) for 
additional information. 


N 
i) 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Dardanus venosus (H. Milne Edwards, 1848) 


Material.—Stations 2, 3 and 5; Aug, Apr and May; 3 tndividuals. 
Measurements.—One male, 6.4 mm sl; two females, 8.3 and 16.1. 
Habitat.—Found in grassbeds (Williams 1965). 

Remarks.—Bright red transverse bands on walking less and chela make this 
species quite distinctive, even after alcohol preservation. Found from North Car- 
olina to the West Indies, including Surinam, Brazil and Bermuda. Found near- 
shore to 91 m, but has been collected to 366 m. Biffar and Provenzano (1972) 
treat this species in their review of the genus. 


Paguristes limonensis McLaughlin and Provenzano, 1974 


Material.—Stations 1—5; all sampling months; 166 individuals. 

Measurements.—Males, 3.3 to 3.6 mm sl; measured females (all ovigerous), 
3.6 to 5.9. 

Remarks.—Over 77% of these hermit crabs were collected at night. The ma- 
jority of individuals were badly damaged during extraction from the shell. Ovi- 
gerous females were taken in July, August, September, November, January, 
March, April and May. This recently described species ranges from the west 
coast of Florida, Panama and Colombia; to 234 m. See McLaughlin and Provan- 
zano (1974) for a more detailed taxonomic discussion. 


SUPERFAMILY PAGUROIDEA 
Family Paguridae 


Pagurus bonairensis Schmitt, 1936 


Material.—Stations 1—5; all months sampled; 190 individuals. 

Measurements.—Ovigerous females, 3.1 to 4.8 mm sl.; remaining specimens, 
Uf (0) Date 

Habitat.—Common in seagrass (Thalassia) beds (in the northeastern Gulf of 
Mexico, Abele 1970; Greening and Livingston 1982; personal obs.). 

Remarks.—Most individuals collected at station 4; over 26% were collected in 
May night trawls. Ovigerous months included July, August, September, Novem- 
ber, March and May. This abundant species was previously misidentified as P. 
annulipes (see Rouse 1970) by Provenzano (1959). Occurs in Florida, Cuba and 
Puerto Rico, although in the northern part of its range this species is often con- 
fused with P. annulipes (Rouse 1970; Garcia-Gomez 1982). Garcia-Gomez (1982) 
provides further clarification. 


Pagurus brevidactylus (Stimpson, 1858) 


Material.—Stations 1—5; all sampling months; 50 individuals. 

Measurements.—Ovigerous females, 3.9 to 5.4 mm sl; remaining intact speci- 
mens, 3.6 to 4.1. 

Habitat.—Provenzano (1959) suggests that this species is most often taken on 
hard bottoms to 229 m; we collected our samples in sandy-vegetated areas less 
than 2 m. 


VOLUME 96, NUMBER 2 213 


Remarks.—Ovigerous months included August, September and February to 
May. This species has been noted from Bermuda, Florida, the Caribbean and 
northeast Gulf of Mexico; intertidal to 50 m. See McLaughlin (1975) and Lemaitre 
et al. (1982) for a reclarification of this species. 


SUPERFAMILY GALATHEOIDEA 
Family Porcellanidae 


Megalobrachium mortenseni Haig, 1962 


Material.—One individual; all additional data lost. 

Measurements.—Male, 4.0 mm cw. 

Habitat.—In sandy areas with calcareous algae (Brazil), coarse sediments 
(gravel) and among rocks. 

Remarks.—Haig (1962) discussed the close resemblance of M. mortensi to M. 
erosum, the latter occurring in the Gulf of California. Gore and Abele (1976) 
considered the two to be geminate species. Reported from the Virgin Islands 
southward to Brazil; to 27 m. This is a new record for Caribbean Panama. 


Petrolisthes armatus (Gibbes, 1850) 


Material.—Stations 1, 2 and 3; Aug to Mar; 38 individuals. 

Measurements.—Male, 3.8 to 7.4 mm cw; ovigerous females, 3.0 to 6.3; non- 
ovigerous females, 3.1 to 4.3. 

Habitat.—Found in oyster and mussel bars, coral rubble, mangroves and around 
pilings (Haig 1960). Gore and Abele (1976) found this species abundant in rocky 
coralline intertidal on both sides of the Panamanian isthmus. 

Remarks.—Ovigerous females found in all collecting months. Haig (1960) re- 
ports ovigerous females also throughout the year among material examined from 
intertidal and subtidal habitats. This porcellanid crab is reported from Bermuda, 
Georgia and central eastern Gulf coast of Florida to Brazil; western Africa and 
eastern Pacific (Gulf of California to Peru); to 20 m. 


Petrolisthes galanthinus (Bosc, 1802) 


Material.—Stations 1, 2, 3 and 5; July to May; 44 individuals. 

Measurements.—Males, 3.6 to 7.0 mm cw; ovigerous females, 4.4 to 8.8; a 
single non-ovigerous female, 4.1. 

Habitat.—Under rocks, coral rubble and on shell and sand bottoms. Also as- 
sociated with sponges, coral and anenomes (Haig 1960; Gore and Abele 1976). 

Remarks.—Ovigerous females were taken July, August, February and March 
(September and October were the only other months when females were taken 
at all). Haig (1960) found ovigerous females in January, February and March; 
southwest Caribbean, ovigery from January to August (Gore and Abele 1976). 
Species common in the Atlantic from North Carolina to Brazil, including the 
Caribbean area and Surinam. In the Pacific, Haig (1960) reported it only from 
Panama, but Gore and Abele (1976) noted its occurrence from Ecuador and Costa 
Rica also. 


214 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


INFRAORDER BRACHYURA 
SECTION DROMIACEA 
SUPERFAMILY DROMIOIDEA 
Family Dromiidae 


Dromidia antillensis Stimpson, 1858 


Material.—Station 3; Apr; 2 individuals. 

Measurements.—Males (two), 17.8 and 18.8 mm cw. 

Habitat.—Most often on rocky substrates. 

Remarks.—These crabs typically carry sponges or tunicates with their fifth leg, 
modified for grasping the dorsal covering; both specimens had sponges. Found 
from Bermuda and North Carolina to the Gulf of Mexico, the West Indies, Co- 
lombia and Surinam to Brazil; littoral to 300 m. See Powers (1977) for an excellent 
summary of this group. 


SECTION OXYSTOMATA 
SUPERFAMILY LEUCOSIOIDEA 
Family Calappidae 


Calappa angusta H. Milne Edwards, 1880 


Material.—Stations 1 and 2; Aug, Nov, Mar and May; 4 individuals. 

Measurements.—Males (four), 6.3 to 45.1 mm cw (45.1 mm, mature). 

Habitat.—From coral, sand, shell-and gravel substrates. 

Distribution.—From Bermuda and North Carolina to Brazil, including the West 
Indies and the Gulf of Mexico; to 200 m. 


Calappa ocellata Holthuis, 1958 


Material.—Stations 1 and 2; both were lost in transit; 2 individuals. 

Habitat.—Among corals and on sandy bottoms. 

Remarks.—This and the previous calappid species possess a large, toothed 
dactyl used to open gastropod shells (often with hermits inside, Vermeij 1982). 
Shoup (1968) discusses this unique predatory behavior. C. ocellata ranges from 
Bermuda and North Carolina to Brazil; to 52 m, though common in shallower 
waters (see Holthuis 1958). 


Hepatus pudibundis (Herbst, 1785) 


Material.—Station 1; specimens lost; 2 individuals. 

Habitat.—On sand, mud and shelly bottoms (Holthuis 1959). 

Remarks.—The low abundance of this and the previous two calappid species 
is surely a result of their habit of remaining partially buried much of the time 
(Pearse, Humm and Wharton 1942), thus escaping collection by trawling. This 
species has been reported from North Carolina to Brazil, including Louisiana and 
Texas in the Gulf of Mexico and from western Africa; intertidal to 49 m. 


VOLUME 96, NUMBER 2 215 


SECTION OXYRHYNCHA 
SUPERFAMILY MAJOIDEA 
Family Majidae 


Chorinus heros (Herbst, 1790) 


Material.—Stations 1, 2 and 3; May night; 6 individuals. 

Measurements.—One male, 11.3 mm cw; females, 10.7 to 17.0. 

Habitat.—Found on rocks, broken shell, coarse sand, sponges, and sabellarid 
reefs (Gore, pers. comm.). 

Distribution.—From Bermuda and central eastern Florida to Cuba and Brazil; 
shallow to 48 m. 


Macrocoeloma diplacanthum (Stimpson, 1860) 


Material.—Stations 1-5; all collecting months; 33 individuals. 

Measurements.—Males from 7.4 to 31.8 mm cw (including lateral spines); non- 
ovigerous females, 16.9 to 28.7; ovigerous females, 21.8 to 28.4. 

Habitat.—Near shallow reefs and on sandy substrates. 

Remarks.—All individuals were collected in day sampling. Ovigerous females 
found in July, August, November and April. Animals were always encrusted with 
numerous sponges, algae and debris. Rathbun (1925) reported individuals from 
Jamaica and Cuba with rhizocephalan infestations, but none were found infested 
here. Species ranges from Key West, Florida to Colombia, including the Bahamas 
and the Caribbean Sea; in shallow water to 24 m. 


Microphrys bicornutus (Latrielle, 1825) 


Material.—Stations 1—5; all months sampled; 195 individuals. 

Measurements.—Mature males, 5.1 to 23.3 mm cw (including lateral spines); 
immature males, 8.8 to 8.9; mature females, 6.2 to 17.6; immature females, 6.3 
to 8.9; ovigerous females, 11.6 to 21.1. 

Habitat.—On reefs, seagrasses, sponges, and a variety of substrates such as 
shell, rock, sand and mud. 

Remarks.—This majid ranked third in overall species abundance. It was most 
common at stations | and 2. Twenty-six ovigerous females were collected (in all 
months but March). These “‘decorator crabs’’ covered with a variety of algae, 
sponges and other sessile invertebrates. M. bicornutus occurs from North Car- 
olina to Brazil, including the Bahamas, the Florida Keys, St. Croix (pers. obs.), 
Colombia and Bermuda; to 30 m. Powers (1977) provides an excellent review of 
this species. Williams (1965) summarizes its known parasites and Hazlett (1979 
and included references), the behavioral literature on this species. This is one of 
the few species dealt with here, whose biology has been studied in some detail. 


Mithrax (Mithraculus) forceps (A. Milne Edwards, 1875) 


Material.—Stations 1—5; in all sampling months; 88 individuals. 
Measurements.—Males, 4.9 to 21.2 mm cw; ovigerous females (17), 6.8 to 16.4; 
non-ovigerous females, 6.8 to 16.4. 


216 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Habitat.—Under coral rubble, in sponges, Thalassia beds, and several coarse 
substrates (coral, shell, and rock). 

Remarks.—Most (87%) were collected during daytime trawls. Ovigerous fe- 
males were found from July to October, January and March to May (all months 
within which females were collected). Abele (1970) noted that this brick red crab 
commonly preyed on barnacles. Recorded from Bermuda, North Carolina to the 
Gulf of Mexico, Trinidad and Colombia; intertidal to 90 m. See Powers (1977) 
for a brief introduction to this species’ biology. 


Mithrax (Mithraculus) ruber (Stimpson, 1871) 


Material.—Station 4; data lost; 1 individual. 

Habitat.—From sand, coral and mud substrates, with sponges and in seagrass- 
es near reefs. 

Distribution.—Puerto Rico, Cuba, St. Thomas, Barbados, Curacao and Colom- 
bia; to 153 m, though usually shallower. 


Pitho Bell, 1835 


Remarks.—A total of 148 individuals belonging to this genus were collected 
over the course of this study. Over 70% were from May night trawls. In daytime 
sampling, most (70%) individuals at station 1; for night samples, the majority (60%) 
at station 3 (May). No Pitho spp. were taken at station 2 (daytime), yet 19 indi- 
viduals (most P. /herminieri) were collected at night (May). 


The following account by species, for the genus Pitho is tentative. We found, 
and others have noted (Abele 1970 and pers. comm.), a great deal of variation 
(especially with size and sex) regarding the lateral teeth of the carapace. Rath- 
bun’s (1925) key to the species employs these teeth as major diagnostic charac- 
ters. Based on our observations of over 140 individuals, other criteria must be 
used to sort out specimens to species, particularly when several are sympatric. 
Rathbun (1925) has some excellent figures (fig. 116 and 117) of both male abdom- 
inal segments and antennal articles, but caution should be exercised when crabs 
are immature or adults are larger than average-sized. Note: totals below may be 
different than original species counts (Heck 1977); some specimens were lost. 


Pitho aculeata (Gibbes, 1850) 


Material.—Stations 1 and 3; Aug, Sept and Mar to May; 9 individuals (tenta- 
tively identified). 

Measurements.—Mature males, 10.6 to 24.6 mm cw; juvenile males, 12.1 to 
14.4; one ovigerous female, 19.3; mature females (non-ovigerous), 20.6 to 22.4; 
one juvenile female, 12.9. 

Habitat.—Sand, shell and mud bottoms with corals; algae (e.g. Sargassum) 
and seagrass. 

Remarks.—All were collected in daytime sampling. Reported from the Baha- 
mas, Florida (west and Keys) to the West Indies and the northern coast of South 
America; in shallow water. 


VOLUME 96, NUMBER 2 217 


Pitho anisodon (von Martens, 1872) 


Material.—Stations, 14; Sept, Oct and May; 62 individuals originally identi- 
fied. 

Measurements.—Mature males, 18.1 to 25.3 mm cw; one immature male, 12.4 
(May); ovigerous females, 23.7 to 26.8 (all May); one immature female, 10.1 
(May); one non-ovigerous female, 26.3 (May). 

Habitat.—Found in rocks, sand, grassbeds, mud and coral bottoms. 

Remarks.—This species was the most abundant Pitho spp. Collected in greatest 
numbers in September, October and May, with the majority (81%) taken at night 
in May (37 of 50 at station 3 for all night samples). The species occurs in the 
Bahamas, Florida, Cuba, Jamaica and Curag¢ao; to 22 m. 


Pitho lherminieri (Schramm, 1867) 


Material.—Stations 1-4; all months sampled; 59 individuals originally identi- 
fied. 

Measurements.—Mature males, 12.3 to 22.9 mm cw; juvenile males, 10.5 to 
13.5; ovigerous females (two), 16.8 and 18.3 (station 3, May night); non-ovigerous 
females (mature), 18.3 to 24.9; immature females, 9.5 to 9.8 (September and May). 

Habitat.—Found on most coarse bottoms and with seagrasses (Lemaitre 1981); 
infrequently on mud. 

Remarks.—Majority of individuals collected at night in May (63%). Noted from 
North Carolina to west Florida, the West Indies to Brazil; to 221 m, though 
usually to 51 m. 


Pitho quadridentata (Miers, 1879) 


Materials.—Stations 1-4; most in Sept and May; 24 individuals originally iden- 
tified. 

Measurements.—Mature males, 11.5 to 25.0 mm cw; one immature male, 13.0 
(May); 4 ovigerous females (Sept and May night), 20.2 to 26.2; juvenile females, 
9.5 to 9.8 (Sept and May night). 

Remarks.—Most individuals from station 1 (62%). Rathbun (1925) discussed 
its resemblance to P. anisodon. Previously reported only from Jamaica. This 
probably constitutes the first record for Panama and Central America. 


Pitho sexdentata Bell, 1835 


Material.—Station 2 night; May; 2 individuals. 

Remarks.—Two specimens originally recorded by Heck (1977). This species 
has previously been collected in the Pacific only (Rathbun 1925; Garth 1958). 
Unfortunately, these specimens could not be relocated. 


Podochela gracilipes Stimpson, 1871 


Material.—Stations 1, 3 and 5; May; 3 males. 
Measurements.—Males, 3.3 to 4.5 mm cw. 
Habitat.—Associated with rocks, gravel, sand, shell and coral. 


218 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Distribution.—From North Carolina to the Gulf of Mexico, the Caribbean Sea, 
including Colombia, Surinam and Brazil; to 220 m, although we collected from 
depths less than 2 m. 


Podochela riisei Stimpson, 1860 


Material.—Stations | and 3; Aug, Oct and Apr; 4 individuals. 

Measurements.—Males, 4.8 to 6.4 mm cw; a single ovigerous female, 9.7 (Apr). 

Habitat.—Found on most coarse substrates, though often with algae (e.g. Sar- 
sassum) and Thalassia (Wass 1955; Greening and Livingston 1982). 

Remarks.—Abele (1970) found this species often covered with bryozoans. It is 
reported from Bermuda and North Carolina to Mexico and Brazil; shallow to 
90 m. 


Podochela sidneyi Rathbun, 1924 


Material.—Station 1; Aug, Feb and May; 3 badly damaged individuals. 
Distribution.—North Carolina, east and west Florida, Texas, Yucatan Penin- 
sula and Cuba; to 186 m. 


[Note: these 3 Podochela spp. are very similar and the identifications are all 
tentative; dactyls of the last 3 legs missing). Wass (1955) suggests that P. sidneyi’s 
sternal plates are flatter than those of P. riisei. | 


Stenorhynchus seticornis (Herbst, 1788) 


Materials.—Stations 3, 4 and 5; Aug to Nov; 9 individuals. 

Measurements.—Males, 9.8 to 18.7 mm cw; two ovigerous females (Septem- 
ber), 10.0 and 10.7; non-ovigerous females, 7.7 to 9.7. 

Habitats.—On coarse bottoms such as rock, gravel, sand, shell and coral rub- 
ble. 

Remarks.—Found from Bermuda, North Carolina to Brazil, including Colom- 
bia; to 1487 m, though more often in shallower waters. Yang (1967) recognized 
three distinct species in the Atlantic. Previous accounts of S. seticornis from the 
eastern Atlantic (West Africa) are incorrect. Manning and Holthuis (1981:304) 
review the current status of this genus in the Atlantic. Powers (1977) briefly lists 
references on the biology of this interesting crab. See Schnever (1978) for some 
recent work on this species. 


SUPERFAMILY PARTHENOPOIDEA 
Family Parthenopidae 


Heterocrypta granulata (Gibbes, 1850) 


Material.—Station 4; August; one individual. 

Measurements.—Male, 12.8 mm cw. 

Habitat.—Found on shelly, mud, sand, gravel, rock and coral bottoms; also 
collected in Thalassia (Lemaitre 1981). 

Remarks.—This parthenopid crab has been reported from Massachusetts to 
Georgia, the Gulf of Mexico, and the West Indies to Brazil; generally less than 
50 m. Gore and Scotto (1979) provide an excellent review of the family. 


VOLUME 96, NUMBER 2 219 


SECTION BRACHYRHYNCHA 
SUPERFAMILY PORTUNOIDEA 
Family Portunidae 


Callinectes danae Smith, 1869 


Material.—Stations 1—5; all sampling months; 66 individuals. 

Measurements.—Mature males, 58.3 to 105.4 mm cw (including lateral spines); 
juvenile males, 20.4 to 69.4; ovigerous females (two), 76.4 and 99.2; females 
(mature), 61.9 to 103.6; juvenile females, 20.9 to 60.3. 

Habitat.—Common in muddy estuaries, in mangroves, algae, and shelly bot- 
toms; from nearly fresh to full strength seawater (Williams 1974b). 

Remarks.—Ovigerous females taken in August and October. Miles (1951) re- 
ported ovigerous females beginning at 67 mm cw. Rathbun (1930) found C. danae 
along high energy sandy beaches and Park (1969) noted its occurrence on the 
seaward sides of islands in Biscayne Bay, Florida. Over 38% of the crabs main- 
tained a rhizocephalan infection (Table 2). This species noted from Bermuda, 
Central Florida and the Yucatan Peninsula to Brazil; to 75 m. Williams (1974b) 
reviews this genus. 


Cronius tumidulus (Stimpson, 1871) 


Material.—Stations 1, 2, 3 and 4; 93% in May (night); 59 individuals. 

Measurements.—Mature males, 15.0 to 35.2 mm cw (including lateral spines); 
juvenile males, 9.0 to 15.5 (May); two ovigerous females (Station 2 and 3, May 
night); mature females, 16.1 to 34.2 (Sept and May); juvenile females, 9.3 to 14.0 
(May only). 

Habitat.—Coral, sand and rocky bottoms and in Sargassum and Thalassia. 

Remarks.—This species has often been reported from Thalassia beds (Rathbun 
1930; Garth 1978); in fact, Park (1969) found it exclusively there. Several crabs 
had rhizocephalans (see Table 2). Noted from Bermuda, the central east and west 
coasts of Florida, the Florida Keys and the Bahamas; 73 m. 


Lupella forceps (Fabricius, 1793) 


Material.—Station 3; Dec; one individual. 

Measurements.—One immature male, 30.5 mm cw (including lateral spines). 

Habitat.—Common offshore on muddy bottoms and in vegetated areas (Park 
1969). 

Distribution.—Found in the West Indies, Colombia and Surinam; Parks (1969) 
collected one individual in Biscayne Bay, Florida; to 15 m. 


Portunus ordwayi (Stimpson, 1860) 


Material.—Stations 1 and 2; May night; 10 individuals. 

Measurements.—Males, 42.8 to 48.8 mm cw (including lateral spines), oviger- 
ous females (two; station 1 May night), 40.7 and 46.0; females (non-ovigerous), 
40.6 to 45.4. 

Habitat.—On sand, gravel, shell and coral rubble bottoms. 

Distribution.—From Massachusetts, Bermuda and North Carolina to the Gulf 
of Mexico, the Caribbean Sea and southward to Brazil; to 106 m. 


220 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


SUPERFAMILY XANTHOIDEA 
Family Goneplacidae 


Cyrtoplax spinidentata (Benedict, 1892) 


Material.—Stations 1 and 3; Dec, Jan and May; 3 individuals. 

Measurements.—Males (three), 9.9 to 16.8 mm cw. 

Habitat.—With sponges and algae. 

Remarks.—This monospecific genus previously known from the West Indies 
(Jamaica, Puerto Rico, St. Thomas and Trinidad); in shallow water. This consti- 
tutes the first record for Panama and Central America. 


Family Xanthidae 
Panopeus occidentalis Saussure, 1857 


Material.—Stations 1—5; all months sampled; 96 individuals. 

Measurements.—Males, 4.0 to 30.5 mm cw; females, 6.0 to 28.8. 

Habitats.—On sand, shell, rock and gravel bottoms, around sponges, mangrove 
roots, pilings; in Thalassia (and algae) in the Bahamas (Garty 1978) and Colombia 
(Lemaitre 1981). 

Remarks.—Most specimens from station | (65%) in day trawls 88%; see also 
observations by Oliveira (1940). Ovigerous females in July, September, and Oc- 
tober. This abundant xanthid ranges from Bermuda, North Carolina to Brazil 
including the West Indies; intertidal to 18 m. Williams (1965) provides a synopsis 
on the biology of this species. 


Pilumnus dasypodus Kingsley, 1879 


Material.—Station 5; Aug; one individual. 

Measurements.—One ovigerous female, 7.5 mm cw. 

Habitat.—Reported on sponges from pilings, jetties and buoys (Abele 1970). 

Remarks.—Ranges from North Carolina to the Gulf of Mexico, and the West 
Indies to Brazil including Colombia; to 29 m. Williams (1965) has summarized 
the ecology of this species. (Note: two other Pilumnus were originally identified 
[KLH] in Panama as P. reticulatus Stimpson [2 specimens] and P. pannosus 
Rathbun [9 specimens]; however, these were lost in transport). 


Discussion 


Of the 58 decapod species discussed here, five or 8.6% represent new records 
for Caribbean Panama and of these, one (Cyrtoplax spinidentata), represents a 
new record for Central America. Both of the two stomatopod species were pre- 
viously known from Central American waters. No endemics were found. 

Although our collection is only a small percentage of the total decapod fauna 
of Panama (estimated to be 1400 species by Abele [1972]), we believe that we 
have good coverage of the grassbed fauna. Those species missed will be mostly 
infaunal, such as alpheids and stomatopods, or rare species, especially those 
active at night. 

The species treated here show close affinities with the Carolinian fauna treated 
by Williams (1965). For example 34 (59%) of the species collected in Panama 


VOLUME 96, NUMBER 2 174 


Table 2.—Summary of parasitic infestations during 1974-75 sampling period. 


Species infested # of individuals* Pertinent data 
Ambidexter symmetricus 1 1 2, 5.8 mm cl; Station | May 
Callinectes sapidus 17 17 2, 53.9-91.8 mm cw 


Aug, Sept, Oct, Dec, Feb 
Mar, Apr, May; Stations 1, 2, 3, & 5 


Cronius tumidulus 2 136,12, 15.8, 18.2 mm cw 


Aug and May; Stations | and 2 


Microphrys bicornutus 3 26,1 2, 8.3-9.1 mm cw 
Oct, Mar and May; Stations 1, 2 & 3 


Panopeus occidentalis 1 1 2, 8.8 mm cw 
Aug, Station 1 


* Note: Rhizocephalan occurrences for all species except A. symmetricus. One processid with 
bopyrid isopod (branchial). 


grassbeds are also found off the Carolinas (Williams 1965). Thirty-seven (64%) of 
our species are known from Bermuda (Markham and McDermott 1981). Most 
of our species are, however, restricted to tropical, subtropical and warm tem- 
perate waters. This is exemplified by the fact that only two of our species (3.4%) 
show up on Williams (1974a) check list of the decapods of the northeastern United 
States (New Jersey to Maine). In addition 14 species (24%) are shared with the 
results of Heck’s (1979) trawling study in turtlegrass beds of the northwestern 
Gulf of Mexico, while only two species (3.4%) are shared with the decapods 
collected in a similar trawling study in eelgrass beds of the lower Chesapeake 
Bay (Heck and Orth 1980). 


Acknowledgments 


We thank L. G. Abele, R. H. Gore, and F. A. Chace, Jr. for their constructive 
criticisms of earlier drafts of the paper. Species identifications were gladly given 
by Drs. Lawrence G. Abele, Florida State University (Decapods) and Patsy 
McLaughlin, Florida International University (hermit crabs). Support for this 
work was provided by the Department of Biological Science (Florida State Uni- 
versity), the Department of Zoology (University of Maryland), by a Smithsonian 
Predoctoral Fellowship to one of us (KLH) and by the Division of Environmental 
Research, Academy of Natural Sciences of Philadelphia. 


Literature Cited 


Abele, L. G. 1970. The marine decapod (Crustacea of the northwestern Gulf of Mexico.—M.S. 
thesis, Florida State University, Tallahassee, 137 pp. 

———. 1972a. Comparative habitat diversity and faunal relationships between the Pacific and 
Caribbean decapod Crustacea of Panama.—Ph.D. dissertation, University of Miami, Florida 
124 pp. 

1972b. Comparative habitat diversity and faunal relationships between the Pacific and Ca- 
ribbean Panamanian decapod Crustacea: a preliminary report with some remarks on the crus- 
tacean fauna of Panama.—Bulletin of the Biological Society of Washington 2:125-138. 
,and N. Blum. 1977. Ecological aspects of the freshwater decapod crustaceans of the Perlas 
Archipelago, Panama.—Biotropica 9:239-252. 


i) 
Ne 
i) 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


, and B. E. Felgenhauer. 1982. Decapoda.—n: S. P. Parker, ed., Synopsis and classification 

of living organisms, Volume 2, pp. 296-326. McGraw-Hill, New York. 

Berrill, M. 1975. Gregarious behavior of the spiny lobster, Panulirus argus (Crustacea, Decapo- 
da).—Bulletin of Marine Science 25:515—522. 

Biffar, T. A., and A. J. Provenzano, Jr. 1972. Biological results of the University of Miami Deep- 
Sea expeditions. 94. A reexamination of Dardanus venosus (H. Milne Edwards) and D. im- 
perator (Miers), with a description of a new species of Dardanus from the western Atlantic 
(Crustacea, Decapoda, Diogenidae).—Bulletin of Marine Science 22:775-805. 

Bowman, T. E., and L. G. Abele. 1982. Classification of the recent Crustacea.—In: L. G. Abele 
(ed.) The Biology of Crustacea, pp. 1-28, Academic Press, New York. 

Bryce, G. W., Jr. 1961. Larval development of Tozeuma carolinense Kingsley, including ecological 
notes on adults.—M.S. thesis, University of North Carolina, Chapel Hill, North Carolina 59 pp. 

Camp, D. K. 1973. Stomatopod Crustacea.—Memoirs of the Hourglass Cruises, Marine Research 
Laboratory, Florida Department of Natural Resources, St. Petersburg, Florida 3(2), 100 pp. 

Chace, F. A., Jr. 1972. The shrimps of the Smithsonian-Bredin Caribbean Expeditions, with a 
summary of the West Indian shallow-water species (Crustacea: Decapoda: Natantia).—Smith- 
sonianian Contributions to Zoology 98, 179 pp. 

Coutiére, H. 1909. The American species of snapping shrimps of the genus Synalpheus.—Proceed- 
ings of the U.S. National Museum 36: 1-93. 

Davis, G. E. 1971. Aggregations of spiny sea urchins, Diadema antillarum, as shelter for young 
spiny lobsters, Panulirus argus.—Transactions of the American Fisheries Society 100:586— 
587. 

——. 1981. Effects of injuries on spiny lobster, Panulirus argus, and implications for fishery 
management.—Fishery Bulletin 78:979. 

Ewald, J. J. 1969. Observations on the biology of Tozeuwma carolinense (Decapoda, Hippolytidae) 
from Florida, with special reference to larval development.—Bulletin of Marine Science 19: 
510-549. 

Felder, D. L. 1973. An annotated key to crabs and lobsters (Decapoda, Reptantia) from coastal 
waters of the northwestern Gulf of Mexico.—Publication No. LSU-56-73-02 of the Center for 
Wetland Resources, L.S.U., Baton Rouge, Louisiana. 103 pp. 

Garcia-Gomez, J. 1982. The Provenzanoi group of hermit crabs (Crustacea, Decapoda, Paguridae) 
in the western Atlantic Part I. Pagurus maclaughlinae, a new species.—Bulletin of Marine 
Science 32:647-655. 

Garth, J. S. 1948. The Brachyura of the ““Askoy’’ Expedition with remarks on carcinological col- 
lecting in the Panama Bight.—Bulletin of the American Museum of Natural History 92(1): 

1-66. 

. 1958. Brachyura of the Pacific coast of America, Oxyrhyncha.—Allan Hancock Pacific Ex- 
peditions. 21(1):1-499. 

—. 1978. Marine biological investigations in the Bahamas. 19. Decapoda Brachyura.—Sarsia 
63:317-333. 

Glynn, P. W. 1972. Observations on the ecology of the Caribbean and Pacific coasts of Panama. 
Bulletin of the Biological Society of Washington 2: 13-30. 

Gore, R. H. 1981. Three new shrimps, and some interesting new records of decapod Crustacea 
from a deep-water coral reef in the Florida Keys.—Proceedings of the Biological Society of 
Washington 94(1):135-162. 

——. 1982. Porcellanid crabs from the coasts of Mexico and Central America (Crustacea: De- 

capoda: Anomura).—Smithsonian Contributions to Zoology 363:1—34. 

, and L. G. Abele. 1976. Shallow water porcelain crabs from the Pacific and adjacent Carib- 

bean waters (Crustacea: Anomura: Porcellanidae).—Smithsonian Contributions to Zoology 

237, 30 pp. 

,and L. E. Scotto. 1979. Crabs of the family Parthenopidae (Crustacea: Brachyura: Oxyrhyn- 

cha) with notes on specimens from the Indian River region of Florida.—Memoirs of the Hour- 

glass Cruises, Marine Research Laboratory, Florida Department of Natural Resources, St. 

Petersburg, Florida 3(6):98 pp. 

Greening, H. S., and R. J. Livingston. 1982. Diel variation in the structure of seagrass-associated 
epibenthic macroinvertebrate communities.—Marine Ecology Progress Series 7:137—156. 

Haig, J. 1960. The Porcellanidae (Crustacea, Anomura) of the eastern Pacific_—Allan Hancock 

Pacific Expeditions 24:1-440. 


VOLUME 96, NUMBER 2 23 


1962. Papers from Dr. Th. Mortensen’s Pacific Expedition 1914-1916 LXXIX: Porcellanid 
crabs from Eastern and Western America.—Videnskabelige Meddelelser fra Dansk Naturhis- 
torisk Forening 1 Kjgbehavn 124:171-192. 

Hazlett, B. A. 1979. Individual distance in Crustacea, III. The spider crab Microphrys bicornutus.— 
Zeitschrift fur Tierpsychologie 49:65—70. 

Heck, K. L., Jr. 1977. Comparative species richness, composition, and abundance of invertebrates 
in Caribbean seagrass (Thalassia testudinum) meadows (Panama).—Marine Biology 41:335- 
348. 

—. 1979. Some determinants of the composition and abundance of motile macroinvertebrate 

species in tropical and temperate turtlegrass (Thalassia testudinim) meadows.—Journal of 

Biogeography 6:183-200. 

, and G. S. Wetstone. 1977. Habitat complexity and invertebrate species richness and abund- 

ance in tropical seagrass meadows.—Journal of Biogeography 4:135-142. 

, and R. J. Orth. 1980. Structural components of eelgrass (Zostera marina) meadows in the 

lower Chesapeake Bay—decapod Crustacea.—Estuaries 3:289-295. 

Holthuis, L. B. 1958. West Indian crabs of the genus Calappa, with a description of three new 
species.—Studies of the Fauna of Curacao and other Caribbean Islands 8:146—180. 

——. 1949. The caridean Crustacea of the Canary Islands.—Zoologische Mededelingen 30:227— 

DSS: 

1959. The Crustacea Decapoda of Suriname (Dutch Guiana).—Zoologische Verhandelingen 
44:1-296. 

Rhandler, N. A. 1964. Sponge as a shelter for young spiny lobsters.—Transactions of the American 
Fisheries Society 93:204. 

Ledoyer, M. 1959. Les Caridea de la frondaison des herbiers de phanérogames de la région de 
Tuléar.—Recueil des Travaux de la Station Marine d’Endoume (Supplement) 8:63—123. 
Lemaitre, R. 1981. Shallow-water crabs (Decapoda, Brachyura) collected in the southern Caribbean 

near Cartagena, Colombia.—Bulletin of Marine Science 31:234—266. 

, P. A. McLaughlin, and J. Garcia-Gomez. 1982. The Provenzanoi group of hermit crabs 

(Crustacea, Decapoda, Paguridae) in the western Atlantic. Part IV. A review of the group, 

with notes on variations and abnormalities.—Bulletin of Marine Science 32:670-701. 

Lyons, W.G. 1970. Scyllarid lobsters (Crustacea, Decapoda).—Memoirs of the Hourglass Cruises, 
Marine Research Laboratory, Florida Department of Natural Resources, St. Petersburg, Flor- 
ida 1(4):74 pp. 

McLaughlin, P. A. 1975. On the identify of Pagurus brevidactylus (Stimpson) (Decapoda: Paguri- 

dae), with the description of a new species of Pagurus from the western Atlantic.—Bulletin of 

Marine Science 25:359-376. 

,and A. J. Provenzano, Jr. 1974. Hermit crabs of the genus Paguristes (Crustacea: Decapoda: 

Diogenidae) from the western Atlantic. Part. II. Descriptions of six new species.—Bulletin of 

Marine Science 25:359-376. 

Manning, R.B. 1961. A redescription of the Palaemonid shrimp, Leander paulensis Ortmann, based 
on material from Florida.—Bulletin of Marine Science 11:525—-536. 

— —. 1969. Stomatopod Crustacea of the western Atlantic.—Studies in Tropical Oceanography 

8: 1-380. 

, and F. A. Chace, Jr. 1971. Shrmps of the family Processidae (Crustacea, Decapoda, Carid- 

ea) from the northwestern Atlantic.—Smithsonian Contributions to Zoology 89, 41 pp. 

, and L. B. Holthuis. 1981. West African brachyuran crabs (Crustacea: Decapoda).— 

Smithsonian Contributions to Zoology 306, 379 pp. 

Markham, J. C., and J. J. McDermott. 1981. A tabulation of the Crustacea decapoda of Bermuda.— 
Proceedings of the Biological Society of Washington 93:1266—1276. 

Miles, R. M. 1951. An analysis of “‘trashfish’’ of shrimp trawlers operating in Apalachicola Bay 
and adjacent Gulf of Mexico.—M.S. thesis, Florida State University, Tallahassee, 45 pp. 

Oliveira, L. P. H. de. 1940. Observacoes preliminares sobre a biologia dos Crustaceos do genero 
Panopeus Milne Edwards, 1834.—Memorias do Instituto Oswaldo Cruz, Rio de Janeiro 35: 
153-171. 

Park, J. R. 1969. A preliminary study of portunid crabs in Biscayne Bay.—Quarterly Journal of the 
Florida Academy of Science 32:12-20. 

Pearse, A. S., J. J. Humm, and G. W. Wharton. 1942. Ecology of sand beaches at Beaufort, NC.— 

Ecological Monographs 12:135—190. 


224 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Perez Farfante,I. 1969. Western Atlantic shrimps of the genus Penaeus.—Fishery Bulletin 67:461— 
591. 

—. 1971. Western Atlantic shrimps of the genus Metapenaeopsis, with descriptions of three 
new species (Crustacea: Decapoda: Penaeidae).—Smithsonian Contributions to Zoology 79, 
37 pp. 

Powers, L. W. 1977. A catalogue and bibliography to the crabs of the Gulf of Mexico.—University 
of Texas Marine Science Institute, Contribution to Marine Science Supplement No. 20, 190 pp. 

Provenzano, A. J., Jr. 1959. The shallow-water hermit crabs of Florida.—Bulletin of Marine Science 
9:349_420. 

Rathbun, M. J. 1918. The grapsoid crabs of America.—Bulletin of the U.S. National Museum 97: 
1-461. 

—. 1925. The spider crabs of America.—Bulletin of the U.S. National Museum 152:1—613. 

—.. 1930. The candroid crabs of America of the families Euryalidae, Portunidae, Atelecyclidae, 
Cancridae, and Xanthidae.—Bulletin of the U.S. National Museum 152:1—609. 

———. 1937. The oxystomatous and allied crabs of America.—Bulletin of the U.S. National Mu- 
seum 166: 1-278. 

Rouse, W. L. 1970. Littoral Crustacea from southwest Florida.—Quarterly Journal of the Florida 
Academy of Science 32:127—152. 

Schnever, G. 1978. In situ observations on the behaviour of and biology of the tropical spider crab 
Stenorhynchus seticornis Herbst (Crustacea, Decapoda, Brachyura).—Jn: D. S. McLusky and 
A. J. Berry (eds.), Physiology and behaviour of marine organisms, pp. 297-302, Pergammon 
Press, New York. 

Shoup, J. B. 1968. Shell opening by crabs of the genus Calappa.—Science 160:887—-888. 

Vermeij, G. J. 1982. Gastropod shell form, breakage and repair in relation to predation by the crab 
Calappa.—Malacologia 23:1—12. 

Voss, G. L. 1956. Protective coloration and habitat of the shrimp Tozeuwma carolinensis Kingsley, 
(Caridea: Hippolytidae).—Bulletin of Marine Science 6:359-363. 

Wass, M. L. 1955. The decapod crustaceans of Alligator Harbor and adjacent inshore water of 
northwestern Florida.—Quarterly Journal of the Florida Academy of Science 18:129-176. 

Weinstein, M. P., and K. L. Heck, Jr. 1979. Ichthyofauna of seagrass Meadows along the Caribbean 
coast of Panama and in the Gulf of Mexico: composition, structure and community ecology.— 
Marine Biology 50:97—-107. 

Williams, A. B. 1965. Marine decapod crustaceans of the Carolinas.—Fishery Bulletin 65:1—298. 

1974a. Marine flora and fauna of the northeastern United States: Crustacea: Decapoda.— 

NOAA Technical Report NMFS Circ-389, 50 pp. 

——. 1974b. The swimming crabs of the genus Callinectes (Decapoda: Portunidae).—Fishery 
Bulletin 72:685-798. 

Yang, W. T. 1967. A study of zoeal, megalopal and early crab stages of some oxyrhynchous crabs 
(Crustacea: Decapoda).—Ph.D. diss., Univ. of Miami, Florida. 459 pp. 


(LDC) Department of Zoology, University of Maryland, College Park, Mary- 
land 20742; (KLH) Division of Environmental Research, The Academy of Natural 
Sciences of Philadelphia, 19th and the Parkway, Philadelphia, Pennsylvania 19103. 


PROC. BIOL. SOC. WASH. 
96(2), 1983, pp. 225-237 


STUDIES OF NEOTROPICAL CADDISFLIES, XXXIV: 
THE GENUS PLECTROMACRONEMA 
(TRICHOPTERA: HYDROPSYCHIDAE) 


Oliver S. Flint, Jr. 


Abstract.—The new species, P. lisae, is described from Mexico and also re- 
corded from Costa Rica. The distributions of the three known species are re- 
viewed and figures of wings and male genitalia given. The larva and pupa of P. 
lisae are described and figured. They inhabit long, silk and sand tubes attached 
to rocks in the bottom gravel in lotic situations and appear to be strongly pred- 
atory. No traces of any capture net were found. A key is presented to the larvae 
of the six Neotropical Macronematinae genera in which this stage is known. 


Plectromacronema was established in 1906 by Georg Ulmer for the new species 
comptum, taken at Santarem, Brazil. Subsequently the species was recorded from 
Venezuela (Navas 1924), Guyana (Mosely 1931), and Suriname (Flint 1974). Na- 
than Banks (1920) described the new genus and species Podomacronema subfus- 
cum from northeastern Argentina. I synonymized this genus in 1967 with Plec- 
tromacronema, but recognized the distinctness of the two species. The genus has 
contained until now these two species, one from northern South America, the 
other from northeastern Argentina. 

In the summer of 1966, when on a field trip to Mexico and Guatemala, my 
daughter Elizabeth (familiarly Lisa) called my attention one dusk to a swarm of 
large, dark caddisflies active over a pool on a small stream in the coastal moun- 
tains of Chiapas, Mexico. As soon as one was caught, it was obvious to me that 
these must be a species of Plectromacronema, but far from the known range of 
the genus. A good series was obtained on this occasion, and the next summer a 
single example was beaten from a riverside tree in Costa Rica. Attempts on 
several occasions to find the immature stages of the genus at the Chiapas site 
were unsuccessful. 

In May of 1981, Dr. Paul J. Spangler, my wife and I travelled to Mexico, where, 
at the invitation of Dr. Joaquin Bueno S., we helped with a field course in aquatic 
insects, then all travelled into southern Chiapas, collecting there. At the Rio 
Lacanja, 22 km east of Ocosingo on the road to Palenque, we discovered some 
interesting long, silken caddisfly tubes attached to large stones in the river bottom. 
Many of them still contained hydropsychid larvae, and a few mature pupae. That 
night many females (and a teneral male) of the new species of Plectromacronema 
came to our lights. Study of the metamorphotypes in Washington proved that at 
long last the larvae and pupae, previously unknown, of this genus were firmly 
associated with the adults. 


Plectromacronema Ulmer 


The three species of the genus fall into two well defined groups based on leg 
structures. Plectromacronema comptum is alone in having a spur count of 1,4,3 


226 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Figs. 1-3. Wings: 1, Plectromacronema comptum; 2, P. subfuscum; 3, P. lisae. 


(Ulmer claimed a second spur on the foreleg, but I can not see anything macro- 
scopic), and in having the basitarsus of the foreleg elongate and nearly cylindrical. 
The other two species have a spur count of 2,4,4, and the basitarsus of the foreleg 
elongate, but at midlength greatly widened and flattened. However, the general 
appearance and uniformity of the male genitalia in all three species precludes 
the restoration of two genera for the two species groups. 

The three species are easily distinguished by the color patterns of the forewings, 
even though there is considerable variation in the exact numbers and placement 
of the spots. Male genitalia differ very slightly between the three species, with 
the tip of the aedeagus offering the best characters. 


Plectromacronema comptum Ulmer 
Figs. 1, 4 


Plectromacronema comptum Ulmer, 1906:63—65.—Fischer, 1963: 163—164.—Flint, 
1974:114-115; 1978:395, 403. 


This species is easily recognized by the coloration which shows three clusters 
of pale marks on the front margin of the forewing. Posteriad of these pale marks 
are many scattered, small, variable, pale spots. 

Biology.—The immature stages of this species are unknown. My few experi- 
ences with adults, as well as collection records, indicate that the species inhabits 
large rivers. Three collection records (Flint 1974) contain the note ‘‘on water 
surface,’ in the evening, or during rainfall. 


VOLUME 96, NUMBER 2 097 


Figs. 4-6. Male genitalia, lateral: 4, Plectromacronema comptum; 5, P. subfuscum; 6, P. lisae. 


Distribution.—In the literature the species is recorded from Brazil, Guyana, 
Surinam, and Venezuela. The following records either add new countries or major 
political subdivisions. French Guiana, 60 mi up Maroni River, Wm. Schaus, | @. 
Venezuela, Edo. Bolivar, Anacoco (6°5'’N, 61°8’W), 60 m, 10-30 Aug 1979, 3 @ 
(IZAM); Rio Cuyuni, El Dorado, 10 Feb 1976, C.M. & O.S. Flint, Jr., 1 @. 


Plectromacronema subfuscum (Banks) 
IMeS5 LZ. 5 


Plectromacronema comptum Ulmer.—UlImer, 1913:392-393 (Misidentification). 
Podomacronema subfuscum Banks 1920:356.—Fischer, 1963:164. 
Plectromacronema subfuscum (Banks).—Flint, 1967:12. 


In coloration this species has a rather dark anterior margin of the forewing, 
with several distinct, white spots in the apical fourth. The pale marks on the basal 
three-fourths of the wing have a distinctive, blurred appearance. 


228 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Biology.—The immature stages of this species are also unknown. I have taken 
adults at lights near small- to medium-sized watercourses, generally with alter- 
nating riffles and pools. Collections at light are almost exclusively females, rarely 
with teneral males appearing. 

Distribution.—This species is found well to the south of the ranges of the other 
two species. It has only been recorded from the Province of Misiones in Argen- 
tina, where I have also made several collections. The following are new country 
records: Brazil, Edo. Santa Catarina, Nova Teutonia (27°11'S, 52°23’W), 28 Oct— 
2 Nov 1939, F. Plaumann, | 6, 12 2 (MCZ, USNM); same, but 7 Jan 1964, 1 o. 
Uruguay, Dpto. Artigas, San Gregorio, 29 Nov 1959, C. S. Carbonell, 4 gd, 10 9 
(FHCU, USNM). Dpto. Paysandu, Puerto Pepe-Aji, | Dec 1959, C. S. Carbonell, 
11 2 (FHCU, USNM). 


Plectromacronema lisae, new species 
Figs. 3, 6—23 


This species is most closely related to P. subfuscum, with which it shares a 
similar spur count and structure of the foreleg, but from which it is easily distin- 
guished by coloration. The pale spots are clear and distinct, and there is a regular 
series of pale marks along the costal margin which are not aggregated into three 
large marks as they are in P. comptum, nor limited to three large spots in the 
apical fourth as they are in P. subfuscum. Differences in the male genitalia be- 
tween the three species are minute and seem to be primarily in the shape of the 
apex of the aedeagus. In P. comptum the tip is more rounded both above and 
below the stem; in the other species the tip is more enlarged above the stem than 
below. In P. lisae this enlargement is rather abrupt and almost as high as long, 
but in P. subfuscum it is distinctly longer than high. 

Adult.—Length of forewing, d 14-18 mm, @ 13-16 mm. Color fuscous; anten- 
nae basally golden yellow, legs stramineous, basitarsus of foreleg infuscate; fore- 
wing fuscous marked with small hyaline spots, those of costal margin being con- 
siderably enlarged. Fifth abdominal sternum anterolaterally with oval, clear, raised 
boss. Male genitalia: Ninth segment annular, slightly produced dorsomesally. 
Tenth tergum elongate, with membranous areas dorsomesally and laterobasally; 
apices pointed, deeply divided dorsomesally. Clasper elongate, slender, basal 
segment nearly parallel-sided; apical segment terete, clearly separated from basal 
segment. Aedeagus with erect, broad, basal section, stem at right angles to base; 
apex with ventral margin confluent with base of stem, dorsal margin produced 
dorsad, about as high as long; internally with single duct with distinct collar 
subapically, and ending in cylindrical cavity extending inward from center of 
posterior surface. 

Larvae.—Length to 16 mm, width to 3 mm. Sclerites yellow with fuscous 
markings. 

Head.—Elongate, about 1/2 times as long as broad. Color basically yellow, 
infuscate dorsally with distinct, pale, muscle scars. Frontoclypeus with anterior 
margin convex; anterior fourth with small accessory setae, with anterolateral 
palmate hairs. Gena with surface smooth, with few small setae; ventrally lacking 
stridulatory grooves, with row of stout, spicate setae along each side; ventral 
ecdysal line lacking on one side of anterior apotome. Labrum rounded anteriorly, 


VOLUME 96, NUMBER 2 229 


Fig. 7. Plectromacronema lisae, larva, lateral. 


almost semicircular in outline; anterior half hairy with anterior margin very hairy. 
Mandibles with strong mesal teeth and multiple, sharp, apical teeth; no mesal 
brushes. Submentum evenly convex anteriorly; setate laterad. Labium very long, 
slender, sclerotized. 

Thorax.—Pronotum dark mesally with pale muscle scars, paler laterad; meso- 
and metanota darkest mesally, paler laterally. Pronotum with anterior margin 
bearing fringe of slender, short hairs; surfaces of all nota smooth, with scattered 


] 


230 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


WS 


11 


Figs. 8-11. Plectromacronema lisae, larva: 8, head and thoracic nota, dorsal; 9, labrum, dorsal; 
10, mandibles, dorsal; 11, maxillolabium and anterior margin of genae, ventral. 


VOLUME 96, NUMBER 2 231 


Fig. 12. Plectromacronema lisae, schematic gill diagram of lateral aspect of first 8 abdominal 
segments. S = a single, long, central stalk with lateral filaments; D = 2 S-type gills with adjacent 
bases. 


small, decumbent hairs and few erect setae. Prosternite transverse, narrow, with 
dark central mark; meso- and metasterna unornamented. Foretrochantin broad, 
tapering to blunt point apically, with 3 or 4 large, bladelike setae dorsally, arising 
from large bases. All legs nearly equal in length, forefemur distinctly widened. 
Foreleg with inner face of femur, tibia and tarsus with rows of enlarged, bladelike 
setae; ventral margin of femur with fringe of long hairs and row of short, bladelike 
setae. Midleg with inner face of tibia and tarsus with rows of enlarged, bladelike 
setae; ventral margin of femur with fringe of long hairs, dorsal margin with scat- 
tered spinous setae. Hindleg with inner face of tibia and tarsus bearing rows of 
short, bladelike setae; ventral margin of femur with sparse fringe of long hairs; 
ventral margin of trochanter with row of very short, enlarged setae in fringe of 
long hairs. No thoracic gills. 

Abdomen.—Gills consisting of central stem bearing many lateral filaments; 
placed as in Fig. 12; double gills above lateral line often with bases separated, 
very small posteriad and virtually lost in lateral line. Lateral line lacking on 
segments 1-3; very long and dense on segments 4—7, curving ventrad anteriorly 
on segment 8, with complete fringes over dorsum along posterior of segments 4 
and 5. Integument with many, darkened, decumbent setae, and few, scattered, 
erect setae. Segments 2-8 with paired, ventrolateral pockets (generally appearing 
to be slightly invaginated with posterior face partially folded over opening, oc- 
casional examples appear to have pockets everted and almost proleg-like with 
covering of crotchets), bearing dense rows of spines whose tips are hooked an- 
teriad. Sternum 9 bearing pair of ovoid sclerites, with posteriorly directed, blade- 
like setae on basal %4, posterior fourth with long setae; ventrolaterally with long 
setae. Anal gills, if present, not showing. Anal proleg with scattered setae; anal 
brush reduced to 4 long setae; anal claw sharply angled ventrad, ventral sole plate 
with dense brush of dark setae. 

Pupa.—Length to 16 mm, width to 3 mm. 

Head.—Labrum semicircular with rounded basolateral lobes; basolateral lobe 
with 5-6 setae, anterolateral margin with 7-10 setae each side. Mandibles scler- 


232 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


wi 


KO 


=S 
Gao : 


ee \ 
a 


(Fe 
2S 

CESS 
32S 2 
Fa RSD 


Figs. 13-18. Plectromacronema lisae, larva: 13, foreleg, inner face; 14, midleg, inner face; 15, 
hindleg, inner face; 16, prosternite; 17, propleuron and coxa, lateral; 18, abdominal hook-pocket, 
anterior to left. 


otized, elongate, almost parallel-sided, inner margin coarsely serrate; basolateral 
surface with brush of setae. Face with scattered setae; vertex with brush of about 
12 setae; basal antennal segment with brush of about 10 setae. 

Thorax.—Meso- and metanota with 2 pairs of short, submesal setae, one pair 
near anterior margin, other at midlength. Coxae of all legs with apical tuft of dark 


VOLUME 96, NUMBER 2 233 


setae. Midleg with tibia and tarsus flattened and broadened; tarsus with short, 
but dense, lateral fringe. 

Abdomen.—Segments 2-7 bearing lateral gills, those on 4—7 being somewhat 
united basally; with ventral gills on 2-7. Hook plates anteriorly on segments 3— 
8, becoming progressively more erect posteriad; posterior plate on segment 3. 
All segments with few erect setae posteriad; segments 3—7 with band of scattered, 
posteriorly-directed, spinous setae along posterior. Eighth sternum with pair of 
oval sclerites bearing many, posteriorly-directed, bladelike setae. Apical pro- 
cesses elongate, slightly arcuate, rugose, apex broad with thin plate dorsally; 
with scattered setae. 

Material.—Holotype, male: Mexico, Edo. Chiapas, rt. 185 km 35 [12 km north 
Arriaga], 7-8 July 1966, Flint & Ortiz. USNM Type 100590. Paratypes: same 
data, 19 3; Rio Lacanja, 22 km north Ocosingo, 19 May 1981, C.M. & O.S. 
Flint, Jr., | d, 21 2; same, but J. Bueno & H. Velasco, 3 d, 8 2 (IBUNAM). 
Costa Rica, [Pcia. Guanacaste], Rio Corobici, [3.4 mi north] Las Canas, 15 June 
1967, Flint & Ortiz, 1 5. Other: Rio Lacanja site, 20 larvae, 7 pupae. 

Biology.—The three collections of this species all were made at fast-flowing, 
clear, cool streams of apparent good water quality with bottom of sand, gravel, 
large stones and boulders. In size they varied greatly; the stream at the type- 
locality is barely half a meter wide by a few decimeters deep, the Rio Lacanja is 
about 10 meters wide by half a meter deep, and the Rio Corobici perhaps about 
15 meters wide. 

The adult males of the type collection were taken just before dusk, flying in a 
swarm up to a meter or so above the water surface over a small, sand-bottomed 
pool. The Costa Rican male was beaten from a low branch of a riverside tree, 
and the Rio Lacanja adults all came to an ultraviolet light after dark. 

The larvae were found in the Rio Lacanja inhabiting jong silken tubes incor- 
porating sand grains and small pebbles which were partially attached to large 
rocks resting in the bottom sand and gravel. They were most frequently found 
where the current was the strongest, flowing between contiguous rocks in a rather 
coarse sand mixture. The tubes did not extend noticeably above the level of the 
sand, and their posterior ends were usually free of the rock, although for most 
of their length they were attached to the rock. When removed from the water 
many larvae crawled from their tubes, as did some of the mature pupae from the 
bottoms of their tubes. 

In the laboratory, the long tubes proved to be difficult to interpret, there being 
no way to be absolutely certain as to which was anterior or posterior, nor if the 
whole tube was present. The longest tube was 70 mm long and from 5—10 mm in 
diameter. The inner surface was formed of a dense mat of silk to the outside of 
which was cemented sand grains. Some parts were only silken, but gave the 
appearance of having been attached to a smooth object, probably the host rock 
or a large pebble. The inner diameter of the tube appears to be slightly larger 
than the diameter of the larva. Several tubes showed a flared, silk-lined opening 
at one end, which I interpret to be the upper end. In no tube could I find any 
type of structured catching net. The pupae inhabit tubes appearing identical to 
those of the larvae. In only one of five tubes still containing pupae did I find a 
mass of debris behind the pupa with admixed larval sclerites; in all the other 
tubes this region was open and the sclerites gone. Those tubes that appeared 


234 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Figs. 19-22. Plectromacronema lisae: 19, pupal head, anterior; 20, pupal hook plates, with ab- 
dominal segment number and anterior or posterior position; 21, pupal apical appendages, dorsal; 22, 
larval anal proleg, lateral. 


VOLUME 96, NUMBER 2 235 


Fig. 23. Plectromacronema lisae, habitat, Rio Lacanja. 


nearly complete contained the pupa near the posterior end (to judge from the 
orientation of the pupa). There was no solid anterior closure, but the anterior 
portion was loosely clogged with sand and debris, sometimes with a loose mesh 
of silken strands at the bottom of this sand. Below this, there was a long portion 
of the tube, one to three times the length of the pupa, that was clean and in which 
the pupa apparently moved. 

Four larval guts were cut open and the contents examined. One was mostly 
empty, but did contain the recognizable remains of a small larva of a polycen- 
tropodid caddisfly and a chironomid head capsule, plus a quantity of an amorphous 
blackish material. A second was filled with the blackish material, recognizable 
sclerites of small arthropods, and apparently large pieces of arthropod cuticle. 
The third contained only a very small amount of fine blackish matter, and the 
fourth, larger quantities of the same. In conjunction with the strongly spined legs, 
rows of spines on the venter of the head, and sharply pointed mandibles, it seems 
probable that the larvae are strongly predatory, but the brushes on the labrum 
and femora suggest that they may also brush fine particulate organic matter from 


236 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


the substrate. Perhaps they utilize the silken lining of the tube to strain fine 
particulate organic matter from the water, in a manner similar to that of the 
Philopotamidae (Wallace and Malas 1976). 


Neotropical Macronematine Larvae 


With the discovery and description of the larvae of Plectromacronema, only 
the immature stages of Centromacronema, Pseudomacronema, and Neolepto- 
nema remain undescribed of the nine Neotropical genera of Macronematinae. It 
is quite probable that the larvae of Neoleptonema will key to Leptonema, as the 
genera are very similar and probably not distinct. It is more difficult to speculate 
on the appearance of the other two genera. They may well run to couplet 5, or 
Centromacronema may approach the appearance of Macronema in couplet 3. 

In order to aid in the recognition of the larvae of the Neotropical macrone- 
matine genera, I offer the following provisional key which incorporates references 
to recent descriptions and name changes. 


1. Foretrochantin very broad, almost rectangular; forefemur broad and trun- 
cate apically; head and thoracic nota very long and slender (Roback 1966, 


as Hydropsychidaeispsl) ie taser eee Synoestropsis 
— Foretrochantin pointed anteriad; apex of forefemur not broad and trun- 

cate; head and thorax not disproportionately elongate ................ 2 
2. Abdominal sterna 2-7 with paired, ventrolateral pockets bearing recurved 

NOOKS. 5.85805. 4 Ee ee ee Plectromacronema 
= Abdomimnal’stemalacking sichssimuctunes™ a -se eee eee ee eee 3 
3. Anal proleg very long and slender, jointed in the middle (Flint and Bueno 

LOS Dy bd |. nc, a pene ELA Da: co any ana Rdg ais.” TRE Wea 0 Macronema 
— Anal prolegs neither exceedingly long nor jointed .................... 4 
4. Head without a carina on genae (Flint and Wallace 1980) ..... Leptonema 
= JHead with a carimasom genae «2.224.406 4e8ed> aa ooo eee 5 
5. Carinae of genal halves meeting at posterior of frontoclypeus, encircling 

the frontal area of the head (Wiggins 1977, as Macronema) ..... Macrostemum 
— Carinae on genal halves not meeting on midline of head, frontal area of 

head open to the posterior (Flint and Wallace 1980) ......... Blepharopus 

Acknowledgments 


The species is dedicated with pleasure to my daughter, Mrs. Elizabeth A. 
Mattingly, whose sharp eyes were instrumental in the initial discovery of this 
species. The excellent habitus drawings (Figs. 7, 8, 18) were prepared by the 
departmental staff artist, Mr. Young T. Sohn. The wing photographs are the fine 
work of Mr. Victor E. Krantz of the National Museum of Natural History. 

Material was supplied by: Ing. C. S. Carbonell, Facultad de Humanidades y 
Ciencias (Departmento de Artropodos), Universidad de la Republica, Montevi- 
deo, Uruguay (FHCU); Dr. Joaquin Bueno Soria, Instituto de Biologia, Univer- 
sidad Nacional Autonoma de Mexico, Mexico City, Mexico (IBUNAM); Ing. C. 
J. Rosales, Instituto de Zoologia Agricola, Facultad de Agronomia, Maracay, Ven- 
ezuela (IZAM); and Ms. M. Thayer, Museum of Comparative Zoology, Harvard 
University, Cambridge, Massachusetts (MCZ). 


VOLUME 96, NUMBER 2 237 


Literature Cited 


Banks, N. 1920. New neuropteroid insects.—Bulletin of the Museum of Comparative Zoology 64: 
299-362. 

Fischer, F. C. J. 1963. Hydropsychidae, Arctopsychidae.—Trichopterorum Catalogus IV: 1-172. 
Flint, O. S., Jr. 1967. Studies of neotropical caddis flies, V: Types of the species described by 
Banks and Hagen.—-Proceedings of the United States National Museum 123(3619): 1-37. 
—. 1974. Studies of neotropical caddisflies, XV: The Trichoptera of Surinam.—Studies on the 

Fauna of Suriname and Other Guianas 14(55): 1-151. 

———. 1978. Studies of neotropical caddisflies, XXII: Hydropsychidae of the Amazon Basin.— 

Amazoniana 3:373—421. 

, and J. Bueno S. 1982. Studies of neotropical caddisflies, XXXII: The immature stages of 

Macronema variipenne Flint & Bueno, with the division of Macronema by the resurrection 

of Macrostemum (Trichoptera: Hydropsychidae).—Proceedings of the Biological Society of 

Washington, 95:358—370. 

, and J. B. Wallace. 1980. Studies of neotropical caddisflies, XXV: The immature stages of 

Blepharopus diaphanus and Leptonema columbianum (Trichoptera: Hydropsychidae).—Pro- 

ceedings of the Biological Society of Washington 93:178-193. 

Mosely, M. E. 1931. Trichoptera and Ephemeroptera of British Guiana, with special reference to 
the Oxford expedition to British Guiana, 1929.—The Entomologist 64: 169-170. 

Navas, L. 1924. Quelques insectes de 1’ Amérique méridionale.—Annales de la Société Scientifique 
de Bruxelles, Documents et Comptes Rendus 43:249-254. 

Roback, S. S. 1966. The Catherwood Foundation Peruvian-Amazon expedition, XI—The Trichop- 
tera larvae and pupae.—Monographs of the Academy of Natural Sciences of Philadelphia 14: 
235-303. 

Ulmer, G. 1906. Neuer beitrag zur kenntnis aussereuropaeischer Trichopteren.—Notes from the 

Leyden Museum 28: 1-116. 

1913. Verzeichnis der sidamerikanischen Trichopteren, mit Bemerkungen liber einzelne 
Arten.—Deutsche Entomologische Zeitschrift 1913:383—414. 

Wallace, J. B., and D. Malas. 1976. The fine structure of capture nets of larval Philopotamidae 
(Trichoptera), with special emphasis on Dolophilodes distinctus.—Canadian Journal of Zoology 
54: 1788-1802. 

Wiggins, G. B. 1977. Larvae of the North American caddisfly genera (Trichoptera).— University of 
Toronto Press, Toronto and Buffalo. 401 pp. 


Department of Entomology, Smithsonian Institution, Washington, D.C. 20560. 


PROC. BIOL. SOC. WASH. 
96(2), 1983, pp. 238-249 


A REVISION OF THE BOGUEIDAE HARTMAN AND 
FAUCHALD, 1971, AND ITS REDUCTION TO 
BOGUEINAE, A SUBFAMILY OF 
MALDANIDAE (POLYCHAETA) 


Paul S. Wolf 


Abstract.—Boguea enigmatica Hartman, 1945, is redescribed and its known 
range extended. Notes on its larval development are presented. The description 
of Boguella ornata Hartman and Fauchald, 1971, is emended. Based on these 
revisions, the family Bogueidae is reduced to Bogueinae, a subfamily of Maldan- 
idae based on morphological and ontogenetic criteria. Keys to the subfamilies of 
Maldanidae, modified from Fauchald (1977), and to the genera of Bogueinae are 
given. 


Hartman (1945) first described Boguea enigmatica from Bogue Sound, North 
Carolina, as a member of the Owentidae. Hartman and Fauchald (1971) later 
described Boguella ornata from deep waters off the coast of New England, at 
which time they also erected the family Bogueidae with Boguea as the type-genus. 
Boguea enigmatica and Boguella ornata have not been recorded outside of their 
type-localities. 

Superficially the Bogueidae are unique among the polychaetes in possessing 
avicular (terebelloid) uncini and yet lacking the head modifications typical of other 
families with avicular uncini (e.g., Terebellidae, Ampharetidae, Sabellidae, and 
Pectinariidae). However, the following redescription of Boguea enigmatica and 
emended description of Boguella ornata show that the Bogueidae share many 
morphological and ontogenetic characters with the Maldanidae; therefore, Bo- 
gueidae is reduced to a subfamily of the Maldanidae. 

Due to the apparent disagreements in definitions of life stages (see Bookhout 
and Horn 1949, and Hermans 1979, for example), it is necessary to define the 
criteria used in this paper for separating larvae, juveniles, and adults. Specimens 
are considered larvae if they still contain some yolk material. Depending upon 
the stage of development, larvae may or may not possess rostrate uncini. Larvae 
examined range from 2 to about 16 setigers long. Ciliated bands were not seen 
on any of the larvae examined. Specimens are considered juveniles if they lack 
yolk material, yet still possess the rostrate uncini in the anterior setigers. Juve- 
niles range from about 15-23 setigers. Adults are those individuals lacking rostrate 
uncini (except perhaps in far posterior setigers), and ranging from about 23-30 
setigers. It should be noted that the above life-stage distinctions pertain only to 
Boguea enigmatica as it is described in this paper. 


Bogueinae, new rank 


Owenlidae (in part).—Hartman, 1945:42. 
Bogueidae.—Hartman and Fauchald, 1971:148.—Fauchald, 1977:135.—Hobson 
and Banse, 1981:20. 


VOLUME 96, NUMBER 2 939 


Diagnosis.—Cephalic plaque absent; cephalic keel and ciliated nuchal slits 
present. Segmental collars absent. Notopodial spines may be present in far pos- 
terior setigers. Neurosetae as avicular (terebelloid) uncini, arranged in single or 
double rows. Pygidium simple or adorned with papillae; anus terminal, without 
a ventral valve. 


Boguea Hartman, 1945 


Diagnosis.—Cephalic keel distinct. Body composed of three well defined re- 
gions. Uncini from setiger 5, in single rows to setiger 9, then in double rows 
except in far posterior setigers. Notopodial spines present in far posterior setigers. 
Pygidium simple, without papillae. 


Boguea enigmatica Hartman, 1945 


Boguea enigmatica Hartman, 1945:42, pl. 7, figs. 4-6.—Hartman and Fauchald, 
1971:148, pl. 23, figs. h-. 


Material examined.—NORTH CAROLINA: Bogue Sound, Summer, 1940, 6- 
10 ft., sand overlaid with shell fragments, holotype (AHF Poly 0391) and 35 
paratypes (AHF Poly 0392); 6 Aug 1962, 2-3 m, sand and shell (primarily Cras- 
sostrea virginica), 15 specimens (ZMC); 10 May 1972, ca 2.5 m, sand and shell, 
2 specimens (both gravid, 1 with brood) (ZMC). SOUTH CAROLINA: Sta. 
0797(4B), 25 Aug 1977, 31°53'N, 80°46’W, 13 m, sand, | juvenile (USNM 059817), 
1 juvenile (Texas Instruments collection). GEORGIA: Sta. 0220(5C), 25 Feb 1977, 
31°08’N, 80°50'W, 19 m, sand, | specimen (Texas Instruments collection); Sta. 
0513(5B), 16 May 1977, 31°12'’N, 81°08’W, 11 m, sand, | specimen (Texas In- 
struments collection); Sta. 0517(5C), 16 May 1977, 31°08'N, 80°50’W, 14 m, sand, 
2 specimens (USNM 059823); 1 juvenile (USNM 059815); 2 specimens (Texas 
Instruments collection); Sta. 0828(5C), 31 Aug 1977, 31°08’N, 80°50’W, 14 m, 
sand, | juvenile (USNM 059816); Sta. 1263(5B), 31°12’N, 81°08’W, 11 m, sand, 
1 specimen (USNM 059822). FLORIDA, Northeast: Sta. 0877(6F), 9 Jan 1977, 
30°23'’N, 80°18’W, 43 m, sand, | juvenile (Texas Instruments collection); Sta. 
0862(6B), 31 Aug 1977, 30°23'’N, 81°15’W, 15 m, sand, 2 specimens (USNM 059820 
and 059821). Tampa Bay: Interstate Electronics Corp. 713TB, Sta. 003, 11 Oct 
1979, 27°37.1'N, 82°54.0'W, 12 m, sand-gravel, 10 specimens (2 females); Sta. 
004, 11 Oct 1979, 27°37.1'N, 82°55.1’W, 10 m, sand, 4 specimens; Sta. 013, 11 
Oct 1979, 27°37.6'N, 82°54.5'’W, sand-gravel, 3 specimens; 723TB, Sta. 001, Jan 
1980, 27°37.6'N, 82°54.5'W, 13 m, sand, | specimen; Sta. 003, Jan 1980, 27°37.1'N, 
82°54.0'W, 12 m, sand-gravel, 3 specimens; Sta. 004, Jan 1980, 27°37.1'N, 
82°55.1’W, 10 m, sand, 1 specimen; Sta. 006, Jan 1980, 27°36.5'N, 82°53.4’W, 12 
m, sand-gravel, 2 specimens. FLORIDA, Southwest: BLM, Sta. 24C, Nov 1980, 
25°16.90'N, 83°43.18’W, 88.4 m, medium sand, | specimen; Sta. 2101H (MAF- 
LA), Feb 1978, 26°24'59.6"N, 82°15'08.9"W, 11 m, sand, | specimen. Pensacola: 
Interstate Electronics Corportion, 732MO, Sta. 012, June 1980, 30°17.2’N, 
87°18.5'W, 6 m, sand, 1 juvenile. ALABAMA: Mobile Bay (Mobil Oil Corpora- 
tion), Sta. 053, July 1978, 30°15'13”"N, 88°03'08”W, 6 m, sand, 1 specimen. MIS- 
SISSIPPI: Horn Island, Corps of Engin-ers, Sta. 023, 7 Nov 1980, 30°15.05'N, 
88°49.78'W, 5.5 m, sand, 33 specimens (including | female with brood of 31 


240 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


larvae); 3 Apr 1981, 44 specimens (including 3 females, | ripe male, 20 juveniles, 
9 larvae). Petit Bois Island, Corps of Engineers, Sta. 038, 22 Oct 1980, 30°13.91'N, 
88°32.48’W, 3.8 m, sand | specimen; 3 Apr 1981, 167 specimens (including 12 
females, 4 ripe males, 39 juveniles, 66 larvae). TEXAS: East Flower Garden Reef, 
U.S. National Marine Fisheries Service, Sta. EFG-IV-6-3, 22 Oct 1981, 27°53.63'N, 
93°39.10’'W, 102 m, silty sand, 2 specimens; Sta. EFG-IV-7-4, 26 Oct 1981, 
27°55.38'N, 93°38.96'W, 101 m, silty sand, | juvenile. 

Redescription.—Length to about 10.0 mm, width to about 0.5 mm. Adults to 
about 30 setigers. Glandular areas present, accentuated by methyl green staining. 
Midventral glandular streak present along entire length of worm from posterior 
rim of mouth. 

Body (Fig. la) divided into 3 regions marked by degree of segmentation, types 
and number of notosetae, number of rows of uncini as well as relative number 
of individual uncini in each row, and development and distribution of glandular 
areas. 

Anterior region.—Comprises prostomium, peristomium, and setigers 1-8. Pro- 
stomium (Fig. la—b) rounded and slightly flattened anteriorly, arched dorsally due 
to presence of cephalic keel; cephalic plaque absent; curved, ciliated nuchal slits 
located laterally. Cilia present around ventral perimeter of prostomium, in addi- 
tion ciliated areas present at least laterally on prostomium and peristomium. 
Exact location and amount of cilia indeterminable due to poor condition of spec- 
imens. 

Prostomium and peristomium fused; peristomium with large rounded mouth 
ventrally. Margin of peristomium and setiger | indistinct except for slight lateral 
indentations. 

Segmentation of anterior region distinct from setiger 2. Setigers 2-8 campan- 
ulate when contracted, each widest anteriorly; when extended, each setiger longer 
than wide, with rounded margins. Collars absent; junction of setigers 8—9 indis- 
tinct. 

Notopodia with simple setae only, emerging from small conical projections near 
middle of each setiger. Setae smooth basally, finely hispid along most of length 
(Fig. Ic). Up to 8 setae per fascicle arranged in 2 rows of 4 each; occasionally 
with 1—2 additional small, thin capillary setae. 

Neuropodia present from setiger 5 as small, slightly elevated tori, each with 
single row of avicular uncini. Each uncinus with about 15 teeth arranged in 3 
crescentic rows surmounting large, anteriorly directed main fang (Fig. Id-—e). 
Uncini numbering 5-9 on setiger 5, gradually increasing to 7-11 on setiger 8. 

Glandular areas of anterior region are shown in Fig. la, after staining with 
methyl green (Banse 1970). Prostomium and peristomium with scattered cells, 
saddled area at cephalic keel, and narrow belt at frontal margin of prostomium. 
Glandular cells of setiger | scattered throughout with additional encircling band 
of cells anterior to notopodia. This band is interrupted by the midventral glandular 
streak. Glandular areas of setigers 2-8 completely encircle the body. 

Middle region.—Comprises setigers 9-14. Segmentation indistinct; setigers 
longer than wide; notopodia and neuropodia as in anterior region. 

Notosetae (Fig. If) similar to those in anterior region but about half as wide 
and numbering only 4 per fascicle arranged in 2 rows of 2 each. 

Uncini in double rows from setiger 9. Anterior row with 2—S uncini, their main 


VOLUME 96, NUMBER 2 


241 


mm 


0.| 


E 
E 
t+ 
° 
co) 


Be 
=e 


= 


ss 
VUABRBSBVSSO TT 


To 


SEN 


0.02mm 


0.0lmm 


0.O0lmm 
(q>) 
0.04mm 
=> 


Fig. 1. Boguea enigmatica: a, Whole worm showing staining pattern, lateral view; b, Prostomium, 
lateral view; c, Hispid notoseta from setiger 2; d, Avicular uncinus from setiger 7, lateral view; e, 
Same, edge-on view; f, Notoseta from setiger 11; g, Channelled notoseta from setiger 16; h, Same, 


diagrammatic cross section; i, Serrate notoseta from setiger 16; j, Notopodial spine from setiger 16; 
k, Juvenile rostrate uncinus from setiger 2. 


242 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


fangs directed posteriorly. Posterior row with 8-14 uncini, their main fangs di- 
rected anteriorly. About 4 uncini added to anterior row by setiger 14 while number 
of uncini in posterior row remains constant or randomly varies by 1-2. Shape 
and dentition of uncini as in anterior region. 

Glandular regions (after methyl green staining) restricted to small areas around 
notopodia and posterior to neuropodial tori; some specimens may also have scat- 
tered cells dorsally on setiger 9 (Fig. la). 

Posterior region.—Comprises remainder of body from setiger 15 to pygidium. 
Segmentation indistinct (except in far posterior setigers) (Fig. la). Setigers grad- 
ually decreasing in length and width posteriorly; noto- and neuropodia as in pre- 
vious body regions. 

Notopodia each with 2 rows of simple setae: anterior row of about 6 setae 
which, viewed laterally, appear unilimbate (Fig. 1g); edge-on view reveals 2 mar- 
gins forming U-shaped channel (Fig. lh). Posterior row of about 6 serrate simple 
setae (Fig. 11), each alternating with channelled setae and strongly curved towards 
anterior end of worm. Notosetae decreasing in number and gradually replaced 
by narrow capillary setae and 1-4 acicular spines (Fig. 1j) in 3—7 posteriormost 
setigers. 

At setiger 15, number of uncini in posterior row abruptly decreases (x = 4.8, 
n = 6); anterior row retains nearly same number of uncini as in preceding setigers. 
Farther back number of uncini in both rows gradually decreases with anterior 
row decreasing more rapidly than posterior row; last 1-2 setigers sometimes 
devoid of uncini. 

Glandular areas of some specimens located only at neuropodial tori. In others, 
even from same locality, cells also present at notopodia and coalesce across 
dorsum. 

Pygidium simple with terminal anus; ventral valve absent. 

Distribution.—Atlantic, North Carolina to northwest Florida; Gulf of Mexico, 
southern Florida to Texas; 2-102 m; primarily sand substrata. 

Notes on larval development.—Boguea enigmatica broods its young. Gravid 
females were found in clear tubes together with eggs and variously developed 
larvae. The broods were found in abandoned serpulid polychaete tubes, but more 
often, the boguein tube was cemented to the concave side of empty bivalve shells 
(e.g., Spisula soldissima and indeterminable venerids in the Mississippi Sound 
area, and Crassostrea virginica in Bogue Sound, North Carolina). Developing 
larvae were found between the body of the mother and the interior wall of the 
tube. Other eggs remained inside the mother’s body. The brood sometimes re- 
vealed all stages of larval development from egg to advanced larval stage (14-16 
setigers). Eggs of one female B. enigmatica measured about 220 wm. There were 
approximately 75 eggs within the female’s body cavity. 

Of the setigerous larvae examined, none has less than 2 pairs of notosetal 
bundles. It appears then, following the reasoning of Bookhout and Horn (1949), 
that the first 2 pairs of bundles appear simultaneously even though the notopodia 
of the first setiger may each contain 2 notosetae while the notopodia of the second 
setiger each contain only | seta. All notosetae are very thin, without discernible 
limbate margins or pinnae as in the adult. The mouth begins forming at the 
3-setiger stage. Rostrate uncini (Fig. 1k) begin developing at the 4-setiger stage 
(Fig. 2a). At the 6-setiger stage (Fig. 2b), nuchal slits are apparent for the first 
time although they do not appear ciliated. 


VOLUME 96, NUMBER 2 743 


noS 


a | nes 


O.1Omm 


C 


Fig. 2. Boguea enigmatica larvae: a, 4-setiger stage; b, 6-setiger stage; c, 10-setiger stage. nS, 
nuchal slit; neS, neuroseta; noS, notosetae; stippling indicates yolk. 


244 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


At the 10-setiger stage (Fig. 2c), a pharynx is visible, segmentation of setigers 
1-5 is distinct, yolk remains posterior to setiger 6, and rostrate uncini are well 
developed anteriorly. Also at the 10-setiger stage, the anus becomes apparent. 

At the 12-setiger stage, the first avicular uncinus appears, developing below 
the rostrate uncinus of setiger 5. At the 15- to 16-setiger stage, the anterior row 
of avicular uncini on setiger 9 is developing and notopodial spines are present in 
the last 1-2 setigers. The adult distribution and types of notosetae are still not 
apparent. All yolk material has been absorbed and segmentation is distinct 
throughout. Fecal pellets are present in some specimens. 

The smallest specimen of B. enigmatica not found with other larvae is 16 
setigers long. This is about the stage at which larvae emerge from the parent 
tube. 

At the 17-setiger stage, some anterior notosetae are limbate and have distal 
pinnae as in the adult. At the 19-setiger stage, the channelled notosetae as well 
as the serrate notosetae of the posterior region are seen developing for the first 
time. 

In general, development of all morphological characters proceeds from the 
anterior to the posterior end. The development of notosetae always precedes that 
of the neurosetae. In contrast, at the 1 1-setiger stage of Axiothella mucosa, uncini 
develop before the notosetae (Bookhout and Horn 1949). 

Juvenile specimens (about 16—23 setigers) are morphologically identical to adults 
except that the juveniles have 2 types of neurosetae. The rostrate uncini (Fig. 
lk) are present, | per fascicle, from setiger 1. Beginning at setiger 5, the rostrate 
uncinus is always located in the superior part of the neuropodial fascicle. As the 
worm matures, the rostrate uncini are lost randomly along the body, usually 
occurring first in the anterior setigers. The specimens are thus left in the typical 
adult condition of having only avicular uncini, except in some far posterior se- 
tigers where occasionally a rostrate uncinus is present. 

The loss of the rostrate uncini gives a clue to setal formation and migration, 
i.e., setae form ventrally in the fascicle and migrate dorsally where they are lost. 
Further evidence for this process is found in observing where the smallest avicular 
uncinus occurs in the fascicie. In developing worms, the smallest uncinus is 
always located dorsally with consecutively larger uncini being formed ventrally. 
This sequence is consistent with that found by Pilgrim (1977) for Clymenella 
torquata (Leidy, 1855) and Euclymene modesta (Quatrefages, 1865). Bookhout 
and Horn (1949, fig. 11) show a rostrate uncinus for larval Axiothella mucosa 
that is quite different from the adult uncinus (Andrews 1891, fig. 35). Day (1967: 
616) states that the first uncini in maldanids appear as ‘‘S-shaped hooks,” al- 
though he does not give specific examples or pertinent citations. Larval uncini 
are reported for members of other families as well, e.g., Sabellidae (Euchone) by 
Banse (1970) and Ampharetidae (Hobsonia florida) by Zottoli (1974). 

That larval uncini can be lost and not replaced, as is the case in the first four 
setigers of B. enigmatica, may not be uncommon. I have observed that juvenile 
specimens of Asychis elongata (Verrill, 1873) collected in Mississippi Sound have 
a pair of rostrate uncini in setiger 1. As the worm matures, the uncini are lost 
leaving the adult arrangement, i.e., uncini absent in setiger 1. 

Remarks.—The staining technique used here was described by Banse (1970). 
The stain delineates, presumably, the mucus-secreting cells. Before staining, 


VOLUME 96, NUMBER 2 345 


glandular cells appear only as thickened, granular regions limited to setigers 2— 
8. Methyl green staining reveals glandular cells as described above and shown in 
Figure la. Larvae and juveniles were not stained. 

This redescription of Boguea enigmatica differs considerably from the original. 
The most noteworthy discrepancy is that uncini begin on setiger 5 instead of 4 
as stated by Hartman (1945). Among the paratypes and other specimens examined 
are individuals with uncini beginning on setiger 4. This is, however, due to re- 
generation of the anterior end. 

As in other maldanids, Boguea enigmatica has a large, sac-like proboscis in- 
dicating that it is a deposit feeder. Cursory gut analysis revealed tests of diatoms 
and protozoans such as Radiolaria. 


Boguella Hartman and Fauchald, 1971, emended 


Diagnosis.—Cephalic keel and nuchal slits poorly defined. Uncini from setiger 
4, in double rows except in far posterior setigers. Both Rhodine-type and avicular 
(terebelloid) uncini present in most setigers. Plumose notosetae present medially. 
Notopodial spines absent. Pygidium papillate. 


Boguella ornata Hartman and Fauchald, 1971, emended 
Boguella ornata Hartman and Fauchald, 1971:149, pl. 23, figs. a—g. 


Material examined. —NEW ENGLAND: Bermuda rise, Sta. A119, 19 Aug 
1966, 32°15.8' to 32°16.1'N, 64°31.6’ to 64°32.6’W, 2095-2223 m, pteropod ooze, 
holotype (AHF Poly 0927), and 16 paratypes (AHF Poly 0928). 

Description.—Length to about 7.0 mm, width to about 0.6 mm. Largest spec- 
imen (holotype) with 25 setigers. Prostomium globular and rounded anteriorly. 
Cephalic keel and nuchal slits poorly defined (Fig. 3a). Prostomium and peri- 
stomium fused; peristomium with small crescentic mouth. Entire head granular 
in appearance (Fig. 3a). Peristomium followed by I—2 long asetigerous segments. 

Segmentation distinct throughout; anterior margins of some anterior setigers 
sometimes collar-like depending upon state of contraction. 

Notosetae of setigers 1-6 of 2 types (Fig. 3b) arranged in 2 rows per fascicle. 
Setae of anterior row stout, abruptly tapering, pubescent; setae of posterior row 
smooth basally, with fine pinnae distally. Notosetae of setigers 7 or 8 through 
15—17 include thin, minutely hispid simple setae and 1-4 long, plumose setae (Fig. 
3c); thereafter notosetae as in setigers 1-6. 

Uncini avicular, beginning on setiger 4 in double rows (Fig. 3d); anterior row 
of 22-setiger paratype composed of smaller, Rhodine-type uncini (Fig. 3e), pos- 
terior row composed of larger terebelloid uncini (Fig. 3f). Uncini of both anterior 
and posterior rows similar in size and terebelloid in shape on setiger 11. Anterior 
row with larger, terebelloid uncini and posterior row with smaller, Rhodine-type 
uncini from setiger 12 to about 17. Posterior row of uncini absent from setiger 
18; all uncini absent from setiger 19. Pygidium as described by Hartman and 
Fauchald (1971:151, pl. 23, fig. c). 

Remarks.—The emended description differs considerably from the original giv- 
en by Hartman and Fauchald (1971:149). One noteworthy discrepancy is the 
setiger on which the uncini are first present. On the holotype and 14 of the 16 


246 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


anR : posR 


1.Omm 
0.02mm 


0.02 mm 


0.0lmm 


0.020mm 


—h 


Fig. 3. Boguella ornata: a, Anterior end, lateral view; b, Notosetae from setiger 4; c, Part of 
plumose notoseta from setiger 12; d, Neuropodial uncini from setiger 4; e, Rhodine-type uncinus 
from setiger 9, lateral view; f, Avicular uncinus from same, lateral view. anR, anterior row; poR, 
posterior row. 


paratypes, the uncini begin on setiger 4; however, on two of the paratypes, the 
uncini begin on setiger 5. On those two specimens, another setiger has been added 
anteriorly, thereby not affecting setal arrangements of posterior setigers, e.g., 
plumose setae still beginning on setiger 7. No specimens contained anterior se- 
tigers with just one row of uncini as originally described. Because of the poor 
condition of all type-material, the distribution of glandular areas could not be 
determined with certainty. It does appear, however, that the heaviest concentra- 
tion of gland cells is in setigers 1-8; in some specimens these areas retained a 
brown pigment. In addition, the prostomium may be completely covered with 
gland cells due to the dense granular appearance of the epidermis (Fig. 3a). In 
some specimens, dorsolateral nuchal slits could be seen under low magnification; 
however, these structures could not be seen on slide-mounted specimens. 

Boguella ornata is known only from its original discovery (Hartman and Fau- 
chald 1971:151). 

Discussion.—During examination of specimens of Boguea enigmatica, includ- 
ing type-material, it became apparent that this species has many characters in 
common with the Maldanidae. These are: 1) presence of a cephalic keel as in the 
Lumbriclymeninae and Rhodininae; 2) presence of ciliated nuchal slits apparently 
identical to those of the Lumbriclymeninae; 3) presence of a large ventral mouth 


VOLUME 96, NUMBER 2 247 


cavity; 4) fused prostomium and peristomium; 5) distinctly segmented anterior 
body region of 8 setigers; 6) fused junction of setigers 8 and 9; 7) indistinctly 
segmented posterior body region; 8) unadorned pygidium as in the Lumbricly- 
meninae and Rhodininae; 9) presence of capillary notosetae arranged in 2 rows 
per fascicle as described for Euclymene oerstedi and Clymenella torquata (Pilgrim 
1977); 10) presence of fine pinnae on the notosetae; 11) change in the type of 
notosetae posteriorly; 12) presence of distinct glandular regions, which are more 
pronounced anteriorly and diffuse posteriorly; 13) presence of uncini in double 
rows, facing in opposite directions in some setigers, as in the Rhodininae; 14) 
uncini beginning on setiger 5 as in some species of Rhodininae; 15) presence of 
rostrate larval uncini that are very similar to typical maldanid uncini and to ros- 
trate uncini of larval Axiothella mucosa (Bookhout and Horn 1949); 16) ventral 
formation and dorsal migration and loss of noto- and neurosetae as in Euclymene 
oerstedi and Clymenella torquata. 

Boguella ornata, in addition to most of the characters above, shares at least 
two additional characters with the Maldanidae: plumose setae, similar to those 
shown for Asychis elongata by Light (1974, fig. 1c); and Rhodine-type uncini. 

Boguea enigmatica differs from maldanids, as currently described, in having: 
1) three constant and well defined body regions; 2) notopodial spines in posterior 
setigers; 3) true avicular uncini; and 4) uncini beginning in single rows, changing 
to double rows, and then back to single rows posteriorly. Boguella ornata ad- 
ditionally differs from maldanids in having a unique papillated pygidium, and 
abruptly tapering, pubescent notosetae. 

Some maldanids may actually have three well defined body regions. Most pres- 
ent descriptions allude to the presence of at least two body regions based pri- 
marily on degree of segmentation. Pilgrim (1977) described three body regions 
for Euclymene oerstedi and Clymenella torquata based on changes and patterns 
in the relative numbers of neurosetae from one region to the next. The body 
regions would be defined differently in these two species if degree of segmentation 
was used. Maldanid body regions cannot be defined until very detailed morpho- 
logical comparisons are made. Included in this review should be a variety of 
morphological features such as the degree of segmentation; the number, type, 
and distribution of notosetae and neurosetae; the distribution of glandular regions; 
and perhaps even certain internal structures such as the alimentary canal and 
nephridia (Pilgrim 1965, 1977). 

Notopodial spines are presently unknown among the maldanids. True avicular 
uncini like those of terebellids are also absent among maldanids; however, the 
uncinus of the Rhodininae resembles the avicular uncini. The Rhodine-type un- 
cinus has a prominent posterior process which is lacking in the avicular uncinus. 
It is particularly noteworthy that Boguella ornata has both types of uncini. Bo- 
guella, then, may be the phylogenetic “‘link’’ between the Rhodininae and newly 
proposed Bogueinae. 

In considering the above discussion, it is concluded that Boguea enigmatica 
and Boguella ornata possess characters which would place them in the Maldan- 
idae. However, no existing subfamily can contain them primarily because of the 
type and distribution of their uncini. It is for this reason that I propose to reduce 
the family Bogueidae to the subfamily Bogueinae within the Maldanidae. 


248 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Key to the Subfamilies of Maldanidae 
(Modified from Fauchald 1977) 


le, Bothicephalicrand analiplaguessalb Senter ernst ce ae 2 
=, Atleastanal’ plague: PReSeMty 6 ke. W hance ke Nee ees ied 9 cake 4 
2, AN ACUI Aes CHING PRC SCI. oc Meg ce eral agave entender ou eer Bogueinae 

Avicular uncini absent, only rostrate uncini present ................ swale 3) 
3. Rostrate uncini in double rows, posterior segments with encircling collars 

ligaice RAR ee, Maca) OM ER Da ade BU n 8, cael aeons eet ee Rhodininae 
— Rostrate uncini in single rows, posterior segments not collared ........ 

Ee ee Pen eee a Aes ASUS (05S RSS lal is Lumbriclymeninae 
4. Cephalic plaque absent, anal plaque present ............... Nicomachinae 
= Both cephalic and anal plaques) presemt ear.) see. 2451. eee 5 
Se AMS GOES a ec aot Ae ral eR ere oie Sy CR TO a ae Maldaninae 
ei AMS COTM i) ob cn Sst ae Ae rok ae RR int ee Euclymeninae 


Key to the Bogueinae Genera 


1. Uncini present from setiger 4, plumose notosetae (Fig. 3c) present in 
Middle :SetuSersy os kee, cee ety 2 SOM a oe to Boguella 
— Uncini present from setiger 5, plumose notosetae absent .......... Boguea 


Acknowledgments 


The author wishes to thank the following people for loan of material: Dr. Mer- 
edith Jones, Smithsonian Institution (USNM); Dr. Mary E. Petersen, Zoological 
Museum, Copenhagen (ZMC); Ms. Susan Williams, Allan Hancock Foundation 
(AHF); and Mr. Peter Miller, Texas Instruments, Inc. The following people re- 
viewed all or parts of the manuscript and offered much appreciated advice and 
encouragement: Drs. Mary E. Petersen (ZMC); Kristian Fauchald (USNM); Wil- 
liam Light, University of Wisconsin—Madison; and Kevin Eckelbarger, Harbor 
Branch Foundation. 

Colleagues at Barry A. Vittor and Associates, Inc., contributed greatly to the 
completion of this paper: Messrs. Michael Ewing, Gary Goeke, Jerry Gathof, and 
Ms. Joan Uebelacker assisted through review and valuable discussions; Ms. Dor- 
othy Cotton typed correspondence and parts of the manuscript; and Mr. Ben 
Jordan prepared the final figures for publication. 

Much of the material examined was collected during work performed under 
contract numbers AA550-CT7-2 (U.S. Bureau of Land Management to Texas 
Instruments, Inc.), 68-01-4610 (Environmental Protection Agency to Interstate 
Electronics Corp.), NA80-GA-C-0075 Work Unit B-2 (U.S. Department of Com- 
merce, National Marine Fisheries Service to TerEco, Inc.), and a contract from 
Mobil Oil Corp. to TechCon, Inc. 

Drs. Andrew Lissner (Interstate Electronics Corp.) and E. A. Kennedy (TerEco, 
Inc.) provided collections for some material examined. 

Barry A. Vittor and Associates, Inc., provided material and partial monetary 
support for this research. 


VOLUME 96, NUMBER 2 749 


Literature Cited 


Andrews, E. A. 1891. Report upon the Annelida Polychaeta of Beaufort, North Carolina.—Pro- 
ceedings of the United States National Museum 14(852):277-302. 

Banse, K. 1970. The small species of Euchone Malmgren (Sabellidae, Polychaeta).—Proceedings 
of the Biological Society of Washington 83(35):387—408. 

Bookhout, C. G., and E. C. Horn. 1949. The development of Axiothella mucosa (Andrews).— 
Journal of Morphology 84:145-183. 

Day, J. H. 1967. A monograph on the Polychaeta of Southern Africa.—British Museum of Natural 
History Publication 656: 1-878. 

Fauchald, K. 1977. The polychaete worms. Definitions and keys to the orders, families and gen- 
era.—Natural History Museum of Los Angeles County, Science Series 28:1—190. 

Hartman, O. 1945. The marine annelids of North Carolina.—Bulletin of Duke University Marine 

Station 2:1—54. 

, and K. Fauchald. 1971. Deep water benthic polychaetous annelids off New England to 

Bermuda and other North Atlantic areas, Part 2.—Allan Hancock Monographs in Marine 

Biology 6:1—327. 

Hermans, C. O. 1979. Polychaete egg sizes, life histories and phylogeny. Jn: S. E. Stancyk (Ed.), 
Reproductive ecology of marine invertebrates.—The Belle W. Baruch Library in Marine Sci- 
ence 9:1-9. 

Hobson, K. D., and K. Banse. 1981. Sedentariate and archiannelid polychaetes of British Columbia 
and Washington.—Canadian Bulletin of Fisheries and Aquatic Sciences 209: 1-144. 

Leidy, J. 1855. Contributions towards a knowledge of the marine invertebrates of the coasts of 
Rhode Island and New Jersey.—Journal of the Academy of Natural Sciences of Philadelphia 
3:135-158. 

Light, W. J. 1974. Occurrence of the Atlantic maldanid Asychis elongata (Annelida, Polychaeta) 
in San Francisco Bay, with comments on its synonymy.—Proceedings of the Biological Society 
of Washington 87(17): 175-184. 

Pilgrim, M. 1965. The functional anatomy and histology of the alimentary canal of the maldanid 
polychaetes Clymenella torquata and Euclymene oerstedi.—Journal of Zoology 147:387-405. 

1977. The functional morphology and possible taxonomic significance of the parapodia of 
the maldanid polychaetes Clymenella torquata and Euclymene oerstedi.—Journal of Mor- 
phology 152(3):281—302. 

Quatrefages, A. de. 1865. Histoire naturelle des Annéles marina et d’eau douce. Annélides et 
Gephyriens 1:1—588. Paris: Librarie Encyclopédique de Roret. 

Verrill, A. E. 1873. Report upon the invertebrate animals of Vineyard Sound and the adjacent 
waters, with an account of the physical characters of the region.—U.S. Fish Commission 


Report for 1871—72:295-778. 
Zottoli, R. A. 1974. Reproduction and larval development of the ampharetid polychaete Amphicteis 


floridus.—Transactions of the American Microscopical Society 93(1):78-89. 


Barry A. Vittor and Associates, Inc., 8100 Cottage Hill Rd., Mobile, Alabama 
36609. 


PROC. BIOL. SOC. WASH. 
96(2), 1983, pp. 250-252 


A NEW SPECIES OF CANCELLARIA (MOLLUSCA: 
CANCELLARIIDAE) FROM THE NORTHERN 
GULF OF MEXICO 


Richard E. Petit 


Abstract.—A previously unknown species of Cancellariidae, Cancellaria 
rosewateri, 1s described. This new species, from the northern Gulf of Mexico, is 
unlike any previously known from the Caribbean, and subgeneric placement is 
not possible as there are no closely related species with which it may be grouped. 
Comparisons are made with a species from the Miocene of California and with 
the Indo-Pacific genus Merica. 


During examination of material dredged by the R/V Oregon in 1962, two spec- 
imens of a unique new deep-water cancellariid were discovered. Subgeneric 
placement of this new species, Cancellaria rosewateri, is not attempted as there 
seem to be no closely related species. Comparison with the Indo-Pacific taxon 
Merica is made under the discussion below. The R/V Oregon also obtained at a 
nearby station a specimen of Gerdiella cingulata Olsson & Bayer, 1972. Gerdiella 
is unknown in the Tertiary of the Caribbean and Panamic areas, but is very similar 
to, if not the same as, Mericella Thiele, 1929, from the western Indian Ocean. 


Class Gastropoda 
Order Caenogastropoda 
Superfamily Cancellariacea Ponder 
Family Cancellariidae Forbes & Hanley 
Cancellaria Lamarck 


Cancellaria Lamarck, 1799: 71. 


Type-species.—Voluta reticulata Linné, 1767, by monotypy. 


Cancellaria rosewateri, new species 
Fig. | 


Material examined.—Holotype: Length 29.3 mm, width 18.6 mm, R/V Oregon 
Station 4156, depth, 200 fathoms, 90 miles SSE of Pascagoula, Mississippi, 29°08’N, 
88°18'W, 19 December 1962, USNM 811465. Paratype: Length 30.4 mm, width 
18.8 mm, R/V Oregon Station 4154, 200 fathoms, 95 miles SSE of Mobile, Ala- 
bama, 29°17'N, 87°39’W, 18 December 1962, USNM 811464. 

Shell description.—Nucleus eroded but apparently smooth, naticoid, of about 
14% whorls. Postnuclear whorls, about 5 in number, rounded, with small but 
distinct shoulder behind which lies a sutural channel. Shell thin but solid, slightly 
translucent, with sculpture of evenly spaced smooth spiral cords, about 16 on 
body whorl with about 8 visible on penultimate whorl and spire whorls. Both 
spiral cords and their interspaces devoid of axial sculpture except for fine growth 
lines. Aperture ovate. Columella concave with 3 almost equal folds, anterior one 
forming edge of short, shallow anterior canal. Inside of aperture lirate, lirations 


N 
‘n 
— 


VOLUME 96, NUMBER 2 


Fig. 1. Cancellaria rosewateri: A, B, C, Holotype (USNM 811465), 29.3 x 18.6 mm; D, Paratype 
(USNM 811464), 30.4 x 18.8 mm. 


2S PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


seeming to match external spiral cords, and extending well into aperture. Shell 
horn-colored, suffused with light brown arranged in 2 indistinct spiral bands, the 
widest occupying posterior half of body whorl, and the other the center third of 
anterior half. Periostracum brown, but not describable from remaining traces. 

Type-locality.—200 fathoms, 90 miles SSE of Pascagoula, Mississippi (R/V 
Oregon Station 4156) 

Distribution.—At present known only from the type-locality and nearby R/V 
Oregon Station 4154. 

Etymology.—Named for Dr. Joseph Rosewater, Curator, Division of Mollusks, 
National Museum of Natural History, Smithsonian Institution, in recognition of 
his contributions to malacology and in appreciation of the assistance he has given 
me Over a period of many years. Dr. Rosewater was also the first to recognize 
this species as new and kindly made the specimens available for study and de- 
scription. 

Discussion.—Comparison of Cancellaria rosewateri with Caribbean and Pan- 
amic species is difficult as there are no known species, Recent or Tertiary, which 
are similar. Cancellaria obtusa Deshayes, 1830, of the Panamic-Pacific Province, 
has similar sculpture but has a straighter columella with unequal plications. Can- 
cellaria darwini Petit, 1970, from the Galapagos Islands, has sculpture that is 
primarily spiral but possesses a straight columella. The shape of the aperture and 
the columella of Cancellaria rosewateri closely resemble those features in species 
of the Indo-Pacific genus Merica. Species of Merica also have spiral sculpture, 
but not in the form of smooth bands as on C. rosewateri. Also, the outer lip of 
Merica is markedly prosocline with a distinct stromboid notch. In C. rosewateri 
the outer lip is only slightly prosocline and lacks any evidence of a stromboid 
notch. Cancellaria rosewateri has many characters in common with C. dalli (An- 
derson & Martin, 1914) as figured by Addicott (1970:117, pl. 15, figs. 11-13, 23, 
27), but that species is described as having two columellar folds. Some European 
Tertiary cancellariids have been placed in Merica by various authors but it is 
doubtful that any of these are congeneric with the Indo-Pacific type. 


Literature Cited 
Addicott, W.O. 1970. Miocene gastropods and biostratigraphy of the Kern River Area, California.— 


United States Geological Survey Professional Paper 642:1—174, pls. 1-21. 
Deshayes, G. P. 1830. Encyclopédie Méthodique. Histoire naturelle des vers 2(1): 1-256. Paris. 


Lamarck, J. B. P. A. 1799. Prodrome d’une nouvelle classification des coquilles. . . —Société 
d’Histoire Naturelle de Paris, Mémoires:63—91. 
Linné, C. von. 1767. Systema naturae per regna tria naturae ... . edition duodecimo reformata. 


Vol. 1, pt. 2. Stockholm. . 

Olsson A. A., and F. M. Bayer. 1972. Gerdiella, a new genus of deep-water cancellanids.—Bulletin 
of Marine Science 22(4):875-880. 

Petit, R. E. 1970. Notes on Cancellariidae (Mollusca: Gastropoda)—II.—Tulane Studies in Geology 
and Paleontology 8(2):83—88, pl. 1. 

Ponder, W. F. 1973. The origin and evolution of the Neogastropoda.—Malacologia 12(2):295-338. 

Thiele, J. 1929. Handbuch der systematischen Weichtierkunde.—Jena, Gustav Fischer, 1929-1935. 
1154 pp. 


Research Associate, Department of Invertebrate Zoology, National Museum 
of Natural History, Smithsonian Institution, Washington, D.C. Mailing address: 
P.O. Box 30, North Myrtle Beach, South Carolina 29582. 


PROC. BIOL. SOC. WASH. 
96(2), 1983, pp. 253-269 


A NEW SPECIES OF POLYMORPHIC FISH, 
CICHLASOMA MINCKLEYI, FROM 
CUATRO CIENEGAS, MEXICO 
(TELEOSTEI: CICHLIDAE) 


Irv Kornfield and Jeffrey N. Taylor 


Abstract.—Cichlasoma minckleyi is described from the Cuatro Ciénegas basin, 
Coahuila, Mexico. Discrete morphological variants occurring sympatrically with- 
in this taxon incorporate differences normally separating distinct congeners. Vari- 
ation is partitioned into two non-overlapping body forms (deep-bodied and slen- 
der-bodied) within which occur two distinct pharyngeal morphs (papilliform and 
molariform), each maintained at high frequencies in natural populations. Field 
observations of matings both between morphs and between forms have estab- 
lished conspecificity of morphological variants. The dichotomous intraspecific 
variation presented by C. minckleyi suggests that phenetic characterization alone 
may be insufficient to delineate biological species within some members of the 
Cichlidae. 


An unusual neotropical cichlid has received considerable attention from evo- 
lutionary biologists because of discrete, pronounced morphological and trophic 
variation. The specific status of this fish had been unresolved because the mag- 
nitude of morphological variation suggested the existence of at least three closely 
related taxa (LaBounty 1974), while biochemical analysis strongly supported the 
existence of a single polymorphic species (Sage and Selander 1975). Recent field 
studies (Kornfield et al. 1982) have confirmed the biochemical analyses and es- 
tablished conspecificity among morphological variants. Described here is Cich- 
lasoma minckleyi, a new species of polymorphic cichlid endemic to the Cuatro 
Ciénegas basin, Coahuila, Mexico. 


Methods 


Standard counts and measurements were recorded following the procedures of 
Taylor and Miller (1980). Measurements and qualitative shape descriptions of 
pharyngeal jaws were taken from Barel et al. (1977) with the following additions: 
mid-pharyngeal jaw width (MPW) is the distance between the margins of the 
dentigerous area of the lower pharyngeal jaw at the midpoint of lower pharyngeal 
length parallel to the line of measured lower pharyngeal width; pharyngeal jaw 
horn width (PHW) is the maximum distance across one horn (ramus) of the lower 
pharyngeal jaw. Head width (HW) is the maximum distance between opercula 
measured over the body (with no caliper compression) at the central, fullest 
portion of the operculum. 

Specimens are deposited in the Museum of Zoology, the University of Michigan 
(UMMZ) and the National Museum of Natural History, Washington, D.C. 
(USNM). 


254 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Cichlasoma minckleyi, new species 
Figs. 1-9, Tables 1-3 


Cichlasoma cf. cyanoguttatum Hubbs and Miller, 1965:52. 

Cichlasoma sp. Taylor and Minckley, 1966:19—21 (including radiographs of troph- 
ic morphs).—Taylor, 1966:161—162, pl. 8, fig. 2; pl. 9, figs. 1-2.—Miller, 1968: 
6.—Minckley, 1969:44-45.—Kornfield and Koehn, 1975:427-437.—Sage and 
Selander, 1975:4669-4673.—Hoogerhoud and Barel, 1978:52-56.—Hutchinson, 
1978:180-181, fig. 113.—Kornfield, 1978:336.—Minckley, 1978:394-397, 399— 
401.—Vermelij and Covich, 1978:836—-837.—Deacon et al., 1979:42.—Liem, 1979: 
121.—Thompson, 1979:680-681, fig. 9e.—Graves and Rosenblatt, 1980:243.— 
Turner and Grosse, 1980:259,269.—Jimenez et al., 1981:409-411.—Kornfield 
et al., 1982:658—-664. 

Parapetenia sp. LaBounty, 1974:5, 46, 54-64, 73, 99, figs. 7-9, 11, 15, 23, 25, 33, 
S45 tablessly Ss —o- 

Cichlasoma minckleyi Kornfield, 1981:96-97 (nomen nudum). 


Diagnosis.—A polymorphic species endemic to the Cuatro Ciénegas basin in 
Coahuila, Mexico, placed tentatively in the Parapetenia group (Regan 1906-08) 
and distinguished from other members of the group by the sympatric occurrence 
of individuals of two discretely-variant body forms (deep-bodied and slender- 
bodied) within each of which two distinct pharyngeal morphs (papilliform and 
molariform) are represented. In addition, there are typically XVI, 10—11 dorsal 
ray elements, V, 8—9 anal fin elements, 14 pectoral fin rays, 11—12 gill rakers, and 
29 vertebrae. The caudal peduncle is relatively slender with the length equal to 
or (usually) greater than the least depth. Sexual dichromatism, as developed in 
breeding adults, is uniquely diagnostic for this species: males are dark-green to 
black with the body and fins peppered with pale bluish spots, while females are 
snowy-white with distinctly contrasting black markings along the flank. 

Holotype. —UMMZ 209434, 93.4 mm SL male, deep-bodied form, papilliform 
morph, Posos de la Becerra, 15.7 km by road SSW of Cuatro Ciénegas de Car- 
ranza, Coahuila, Mexico, R. R. Miller and family, C. L. Hubbs, W. L. Minckley, 
D. R. Tindall, and J. E. Craddock, 6 Apr 1961. 

Paratypes.—(all from Coahuila, Mexico): UMMZ 130385 (9), 20-54 mm, and 
UMMZ 130395 (19), 18-70 mm, Cuatro Ciénegas, E. G. Marsh, Jr., 18 Sept 1939; 
UMMZ 179180 (2), 69 and 84 mm, Escobeda, hot spring 10.1 km S of Cuatro 
Ciénegas, W. L. Minckley and J. E. Craddock, 17 Aug 1960; UMMZ 179184 (1), 
28 mm, irrigation ditch (La Angostura) from El] Garabatal River, 2.6 km S of 
Cuatro Ciénegas, Minckley and Craddock, 18 Aug 1960; UMMZ 179194 (148), 14— 
34 mm, ditch (La Angostura canal) from El Garabatal, 6.6 km WSW of Cuatro 
Ciénegas, Minckley, Craddock, and others, 19 Aug 1960; UMMZ 179200 (9), 14— 
74 mm, Laguna San Marcos at San Marcos, 24.2 km by road SSW of Cuatro 
Ciénegas, Minckley, Craddock, and others, 20 Aug 1960; UMMZ 179216 (5), 9- 
144 mm, Rio Mesquites, 9.0 km S of Cuatro Ciénegas, Minckley, Craddock, and 
others, 21-22 Aug 1960; UMMZ 179224 (12), 15-31 mm, Altamira (6.4 km ENE 
of El Candido), 16 km S and 4.8 km E of Cuatro Ciénegas, Minckley and Crad- 
dock, 23 Aug 1960; UMMZ 179832 and USNM 229347 (155), 25-135 mm, col- 
lected with holotype; UMMZ 179838 (25), 28-114 mm, Puente Colorado, 10.2 
km S of Cuatro Ciénegas, Miller, Hubbs, Minckley, and Tindall, 6 Apr 1961; 


255 


VOLUME 96, NUMBER 2 


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256 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 1. Cichlasoma minckleyi, new species, UMMZ 209434, male, 93.4 mm SL, holotype, deep- 
bodied form, papilliform pharyngeal morph, Posos de la Becerra, Cuatro Ciénegas de Carranza, 
Coahuila, Mexico, 1961. 


UMMZ 179859 (74), 25-164 mm, Tierra Blanca, 10.2 km WSW of Cuatro 
Ciénegas, Miller, Hubbs, Minckley, and P. Lugo, 8 Apr 1961; UMMZ 179877 (2), 
20 and 25 mm, Puente Chiquito, 6.7 km S of Cuatro Ciénegas, Miller, Hubbs, 
and Minckley, 10 Apr 1961; UMMZ 179881 (27), 10-91 mm, Laguna Churince, 
21.6 km by road SSW of Cuatro Ciénegas, Miller family, Hubbs, Minckley, and 
Tindall, 10 Apr 1961; UMMZ 180465 (one skeleton), 169 mm, pools along La 
Angostura canal near Garabatal River, ca. 8 km WNW of tip of San Marcos Mt., 
Cuatro Ciénegas basin, W. L. and B. A. Minckley and P. Lugo, 18 Apr 1963; 
UMMZ 198937 (28 preserved and 3 skeletons), 28-149 mm, Rio Mesquites at Los 
Corrales, Cuatro Ciénegas basin, G. R. Smith, J. N. Taylor, and P. Yant, 25 Mar 
1975; UMMZ 198942 (1), 39 mm, small tributary to Rio Mesquites at Los Corrales, 
Cuatro Ciénegas basin, Smith and Yant, 26 Mar 1975; UMMZ 198947 (89 pre- 
served and 36 skeletons), 18—98 mm, lagunas at El Mojarral, 8.7 km SW of Cuatro 
Ciénegas, Smith, Taylor, and Yant, 26 Mar 1975; USNM 231944 (3), 123-140 
mm, lagunas at El Mojarral, I. L. Kornfield and D. C. Smith, 29 July 1979; USNM 
231945 (126), 54-77 mm, Posos de la Becerra, Kornfield and Smith, 12 Aug 1979; 
USNM 231946 (38), 23-51, lagunas at El Mojarral, D. C. Smith, 7-12 July 1980; 
USNM 231947 (2 skeletons), 120 mm, lagunas at El Mojarral, Kornfield, 16 July 
1981. 


VOLUME 96, NUMBER 2 ZT) 


Fig. 2. Cichlasoma minckleyi, new species, UMMZ 179832 (ID #122.1), male, 93.5 mm SL, 
paratype, slender-bodied form, papilliform pharyngeal morph, Posos de la Becerra, Cuatro Ciénegas 
de Carranza, Coahuila, Mexico, 1961. 


Description.—Body forms and color pattern are shown in Figs. 1-2. Propor- 
tional measurements are given in Table 1, and meristic data appear below (no 
significant differences in counts exist between either body forms or pharyngeal 
morphs). 

Dorsal spines, XV(7), XVI(66), XVII(1); dorsal soft rays, 9(3), 10(24), 11(43), 
12(4); anal spines, IV(1), V(68), VI(5); anal soft rays, 7(4), 8(55), 9(15); pectoral 
rays (both fins counted in 25 specimens), 13(3), 14(38), 15(9). 

Scales: lateral series, 28(2), 29(1), 30(5), 31(14), 32(1); upper lateral line, 17(1), 
18(2), 19(7), 20(7), 21(5), 22(1), and lower lateral line, 10(3), 11(6), 12(8), 13(6); 
scale-row overlap of upper lateral line on lower, —1(3), 0(4), 1(9), 2(5), 3(2); 
transverse scales, 17(6), 18(14), 19(2); upper lateral line to soft dorsal origin (not 
including scales on scaly sheath along fin base), 3(20), 314(3); upper lateral line 
to origin of anal fin, 11(20), 12(3); rows around caudal peduncle, 17(1), 18(3), 
19(12), 20(3); rows on cheek, 5(4), 6(14), 7(3). 

Gill rakers on first arch (total including all anterior rudiments): upper limb, 
3(30), 4(40); lower limb, 7(5), 8(46), 9(19); total, 11(25), 12(36), 13(9). 

Vertebrae: precaudal, 13(21), 14(17); caudal, 15(23), 16(15); total, 28(7), 29(30), 
30(1). 

Deep-bodied form (Fig. 1): Body relatively stout, depth 2.2-2.5 in SL; head 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


GY, 


OE ol 
» “ } Py. : Pp ° en" ’ i 


a a care 
BN WU aL 


Fig. 3. Lower pharyngeal jaws defining papilliform pharyngeal morph (above) and molariform 
pharyngeal morph (below). Both specimens 120 mm SL, male, USNM 231947. 


VOLUME 96, NUMBER 2 259 


W £ 
O O 


N 
O 


Mid Pharyngeal Width (MPW) 


© Papilliform 
@ Molariform 


40 60 80 100 120 
Lower Pharyngeal Width (LPW ) 


Fig. 4. Regression of mid-pharyngeal width (MPW) on maximum width of lower pharyngeal (LPW) 
for papilliform and molariform pharyngeal jaw morphs. 


short, length much less than body depth over pelvic base; predorsal contour 
smoothly convex in subadults, becoming straighter and steeper with age (large 
adult males often develop a pronounced nuchal hump, producing a concavity 
over the orbit); prepelvic contour approximately straight, not so steep as pre- 
dorsal; greatest body depth at or near origin of dorsal. Mouth relatively small, 
horizontal to slightly oblique; jaws approximately equal anteriorly. Ascending 
process of premaxillary extending to above anterior margin of orbit. Frenum along 
margin of lower lip generally weak or absent. Gill rakers well-spaced, short and 
stout. . 
Dorsal base of moderate length, originating above or slightly behind insertion 
of pectoral fin; dorsal spines increasing rapidly in length to fifth or sixth, then 
more gradually to last, which is approximately 0.45 head length in individuals 
over 70 mm SL. Soft rays of dorsal fin when relaxed not reaching beyond anterior 
third of caudal fin (except when prolonged in adults). Pectoral fin rounded, asym- 
metric, equal to approximately two-thirds head length, rarely extending to anal 
fin origin. Pelvic fin usually extending to vent or beyond. Caudal fin rounded; 
peduncle length equal to or greater than least depth. 

Dental arcade rounded, jaw teeth unicuspid, widely and regularly set, implanted 
erect in few bands with outer row prominent. Anterior pair in outer row of upper 
jaw largest, conical and acutely pointed, with tips slightly to strongly recurved 


260 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


(PHW) 


Horn Width 


Pharyngeal 


O Papilliform 
@ Molariform 


2O 8 40 60 £80 lOO =6120 
Lower Pharyngeal Width (LPW) 


Fig. 5. Regression of pharyngeal jaw horn width (PHW) on maximum width of lower pharyngeal 
jaw (LPW) for papilliform and molariform pharyngeal jaw morphs. 


(often worn in large specimens), usually lacking posterior cusp. Flanking teeth 
in upper jaw less recurved, gradually decreasing in size laterally. Anterior pair 
of teeth in lower jaw similar in shape but smaller than central pair in upper, 
equivalent to flanking teeth which gradually decrease in size. 

Occlusal surface of lower pharyngeal jaw with dentigerous portion 1.1 to 1.3 
times broader than long (in 20 individuals 51-121 mm SL). Pharyngeal dentition 
dimorphic, with molars in central rows either well-developed (=molariform morph) 
or lacking (=papilliform morph; see Fig. 3). Dentition in papilliform morphs with 
numerous slender unicuspid teeth arranged in irregular rows; dentition in molar- 
iform morphs with 4-8 massive molars in each of two median rows, decreasing 
in size anteriorly, increasing in number with SL, flanked laterally by 1-2 irregular 
rows of smaller molars, with small papilliform teeth distributed irregularly over 
remaining occlusal surface. Lower pharyngeal of papilliform morph with narrower 
mid-occlusal width and narrower rami (Figs. 4—5). 


VOLUME 96, NUMBER 2 261 


Slender-bodied form (Fig. 2): Body elongate, depth 2.6—2.8 in SL (n = 15, 54.3- 
144.7 mm SL); head relatively long, length equal to or greater than body depth 
over pelvic base. Mouth relatively large, oblique; lower jaw projecting beyond 
upper. Caudal peduncle slender, length greater than least depth. 

Coloration.—Basal color (in preservation) variable, ranging from tan to dark 
brown or black; a series of 4—7 black rectangular blotches centered between the 
upper lateral line and anterior extension of the lower lateral line (Fig. 1), the first 
typically originating above midpoint of pectoral fin and the last under the posterior 
soft dorsal. Blotches often overlie fainter crossbars, most conspicuous in sub- 
adults but frequently obscured by dark basal coloration in adult males. Caudal 
spot, centered above lower lateral line at peduncle base, present in subadults, 
but often faint or lacking in dark-colored adults. Adults also occasionally develop 
diffuse black speckling over the flanks and lighter, irregular spotting on the un- 
paired fins. Paired fins dusky or hyaline. 

In life, basal color in nonreproductive individuals varies from light grey or tan 
through yellowish-green to dark green or grey, while markings on the flank are 
black; both ground color and development of flank pattern are subject to rapid 
alterations in particular individuals. Sexual dichromatism in breeding adults is 
marked. Adult males are dark-green to black (often with the flank pattern com- 
pletely obscured) with light blue spots distributed over much of the head, flanks, 
and vertical fins. In spectacular contrast, the ground color in breeding females is 
an intense, snowy white; the black flank markings are conspicuous, but blue 
spotting is absent. 

Comparisons.—Characters that distinguish C. minckleyi from other Cichlaso- 
ma species are summarized in the diagnosis above. Though its origins are ob- 
scure, C. minckleyi appears to be more closely related to endemic cichlids of the 
Rio Panuco drainage, the first major river south of the Rio Grande, than to C. 
cyanoguttatum, which ranges northward into Texas and has recently gained ac- 
cess to the Cuatro Ciénegas basin via newly constructed irrigation systems. This 
hypothesis of relationships is supported by association of both the Panuco species— 
specifically, C. labridens (Pellegrin), C. bartoni (Bean), and C. steindachneri 
Jordan and Snyder—and C. minckleyi with the Parapetenia group within Cich- 
lasoma (Regan 1906—08; LaBounty 1974; Taylor and Miller, in press), while C. 
cyanoguttatum is a member of the Herichthys group (recognized as a genus by 
Regan 1906-08, but placed in Cichlasoma by Meek 1904, and most subsequent 
authors). These two groupings have traditionally been distinguished on the basis 
of jaw dentition; Parapetenia species have fewer, more widely-spaced, conical 
teeth, with one or more anterior pairs in each jaw enlarged to form canines, while 
the teeth in Herichthys species are more numerous, closely-spaced, compressed 
and incisor-like distally. Unfortunately, such a distinction is not as clear-cut as 
originally formulated. Several so-called Parapetenia species, including C. minck- 
leyi and C. labridens, have only weakly enlarged anterior teeth at best, providing 
intermediate dentitional types between Parapetenia and Theraps, a third grouping 
within Cichlasoma characterized by generalized conical jaw teeth but no canines 
(Regan 1906-08). Further, variability in jaw dentition within populations of C. 
cyanoguttatum can be extensive and include relatively noncompressed, conical 
patterns; indeed, C. pavonaceum (Garman), a species recently placed in the 
synonymy of C. cyanoguttatum (Taylor and Miller, in press) was included in the 
Theraps group by Regan (1906-08). Clarification of supraspecific relationships 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 6. Pharyngobranchial apophysis from molariform pharyngeal jaw morph (right) and papilli- 
form jaw morph (left). Both specimens 120 mm SL, male, USNM 231947. 


among these species awaits renewed phylogenetic study of subgroupings within 
the genus Cichlasoma. 

Intraspecific variation observed in Cichlasoma minckleyi mimics morphological 
diversity encountered among other species within Cichlasoma. In C. minckleyi, 
variation related to both pharyngeal architecture and body shape is present. In 


VOLUME 96, NUMBER 2 263 


© Papilliform 


@ Molariform 


Gut Length (cm) 


40 60 80 eye) 120 140 I60 
Standard Length (mm) 


Fig. 7. Regression of gut length on standard length for papilliform and molariform pharyngeal jaw 
morphs. 


the deep-bodied form (Fig. 1), two suites of integrated morphological characters 
accompany the dichotomy in pharyngeal dentition (Table 2). The lower pharyn- 
geal jaw in papilliform morphs is more delicate, possessing a significantly nar- 
rower outline (Fig. 4) with smaller horns (Fig. 5) than the molariform morph. The 
size and position of branchial musculature differs conspicuously between morphs. 
The molariform pharyngeal apparatus is characterized by general hypertrophy 
with marked enlargement of the levator externis, levator posterior and retractor 
dorsalis muscles (Hoogerhoud and Barel 1978). Differences in neurocranial artic- 
ulation of the upper pharyngeal jaw are dramatic, with dorsal support in the 
molariform morph provided by a massive pharyngobranchial apophysis (Fig. 6). 
Correlated with the pharyngeal modifications in deep-bodied forms is a dimor- 
phism in intestinal length; gut length in the molariform morph is significantly less 
than that in the papilliform morph (Fig. 7). A degree of trophic (dietary) special- 
ization occurs between pharyngeal morphs (Taylor and Minckley 1966; LaBounty 
1974; Sage and Selander 1975), but is not as pronounced as originally believed 
(Smith 1982). Gut content analysis indicates that gastropods occur in molariform 
morphs but are usually, though not invariably, absent from guts of papilliform 
morphs. However, gastropods form only a minor component of all food items 
and dietary overlap between morphs is relatively great, particularly in juvenile 
and subadult fishes. 

In addition to these internal characters, pharyngeal morphs of deep-bodied 
forms can be distinguished externally by a significant difference in head width 
(Fig. 8; C. D. N. Barel, pers. comm.) caused by the massiveness of pharyngeal 
and associated musculature in the molariform morph. In the field, this difference 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


264 


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VOLUME 96, NUMBER 2 265 


32 


O Papilliform 


30 @ Molariform 


28 


26 


24 


Head Width (mm) 


(Ze 


20 


44 46 48 50 52 54 56 58 
Head Length (mm) 


Fig. 8. Regression of head width (HW) on standard length for papilliform and molariform pha- 
ryngeal jaw morphs. 


is not apparent to the human observer, but may perhaps be perceptible by the 
fish, particularly since opercular extension is a frequent element of territorial 
behavior in this and allied species (Baerends and Baerends-Van Roon 1950; pers. 
observ.). If perceptible, head width dimorphism could provide a convenient signal 
potentially preadapting pharyngeal morphs for assortative mating. 

Morphological diversity encountered between deep-bodied and slender-bodied 
forms is pronounced. The streamlined form (Fig. 2) is characterized by differences 
in a number of external characters (Table 1) that become more marked with age. 
Though the differences may not be apparent in small fish, no adults with com- 
pletely intermediate morphology have been recognized. Gut analysis of slender- 
bodied forms suggests that they principally consume other fishes (Taylor and 
Minckley 1966; LaBounty 1974; Sage and Selander 1975), a rare food item in the 
diet of deep-bodied forms; correspondingly, gut length in slender-bodied forms 
is usually shorter than that seen in deep-bodied individuals (Table 1). The external 
shape and dietary dichotomy between body forms does not involve pharyngeal 
morphology; both papilliform and molariform dentitions have been observed among 
slender-bodied individuals (LaBounty 1974). 

Sympatric representation of dichotomous morphologies with respect to pha- 
ryngeal dentition and body form has provided a compelling basis to recognize 
multiple taxa within the Cuatro Ciénegas basin (Taylor and Minckley 1966; 


266 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Table 3.—Relative frequency of the molaniform phenotype, by sex, in deep-bodied forms from five 
localities in the Cuatro Ciénegas basin. Sample size in parentheses. 


Laguna Rancho Posos de la Laguna 

El] Mojarral Orozco Escobeda Becerra Churince 
36d 0.583 (26) 0.500 (16) 0.462 (13) 0.384 (86) 0.375 (24) 
QE 0.400 (35) 0.294 (17) 0.111 (9) 0.122 (156) 0.250 (28) 


Minckley 1969; LaBounty 1974; Kornfield and Koehn 1975). The diversity seen 
within C. minckleyi is precisely the kind which characterizes differences among 
distinct biological species within the genus Cichlasoma, as illustrated, for ex- 
ample, by the endemic cichlid fauna of the Rio Panuco basin (Taylor and Miller, 
in press). In the Rio Verde, a sympatric pair of deep-bodied species, Cichlasoma 
bartoni and C. labridens, exhibits a comparable dichotomy in pharyngeal denti- 
tion, while in the Rio Gallinas system, C. labridens and C. steindachneri differ 
in both body form and pharyngeal dentition. Unlike the Cuatro Ciénegas situation, 
however, the Rio Panuco species can also be distinguished on the basis of breed- 
ing coloration, electrophoretic comparisons, and a number of meristic character- 
istics. Nevertheless, conventional morphological treatment of C. minckleyi con- 
cluded that the taxa consisted of at least three species within the Cuatro Ciénegas 
basin. However, biochemical studies demonstrating parallel variation in isozyme 
frequencies between pharyngeal morphs among isolated localities strongly sup- 
ported a single-species hypothesis (Sage and Selander 1975). This systematic 
problem has been resolved by field observations of reproductive behavior. Suc- 
cessful matings both between deep-bodied fish of different pharyngeal morphs 
(Kornfield et al. 1982) and between different body forms (Taylor, unpubl.) con- 
vincingly established conspecificity. As a corollary, this finding emphasizes the 
potential insufficiency of phenetic (=morphological) criteria in delimiting species 
within the family Cichlidae. 

The relative contributions of genetic and environmental factors to total phe- 
notypic variation within C. minckleyi are unclear. While most deep-bodied spec- 
imens can be easily assigned to a specific pharyngeal morph, a small percentage 
of fish (<5%) exhibit intermediate pharyngeal morphologies and gut lengths. The 
genetic basis for intermediacy is unknown, but cannot be due to hybridization in 
the conventional sense. Proportions of the two pharyngeal morphs vary in dif- 
ferent size-classes and also can differ considerably among localities. However, 
within localities, the proportion of individuals with molariform dentition is sub- 
stantially greater in males than in females (Table 3). While such a difference might 
reflect differential selective pressures, its consistency across localities suggests a 
simple model for the inheritance of alternate pharyngeal states. Assuming that 
pharyngeal phenotype is regulated by a single diallelic locus with a recessive 
molariform allele, the difference in the relative proportion of the two types be- 
tween males and females is consistent with sex linkage (Fig. 9). However, ac- 
curate recognition of control over dentition and body form will require further 
studies of formal inheritance. 

Distribution.—Cichlasoma minckleyi is known only from the Cuatro Ciénegas 
basin, Coahuila, Mexico. Within the basin it is common and widely distributed 
in most streams, ponds and lagunas. At a few localities, in the southeastern part 


VOLUME 96, NUMBER 2 267 


> 
S 1.0 
z 

WwW 

S 

a 0.8 
a 

Le 

= 06 
og 

O 

Le 

a 

aq 0.4 
J 

oO 

= 

et 0.2 
—) 

<q 

a 

re O 


O 0.2 0.4 0.6 08 1.0 
MALE MOLARIFORM FREQUENCY 


Fig. 9. Observed relative frequencies of molariform pharyngeal jaw phenotype in males and fe- 
males from five localities in the Cuatro Ciénegas basin. Curve represents expected relationship when 
molariform phenotype is recessive and sex-linked. 


of the basin, including Laguna Santa Tecla, it occurs sympatrically with the recent 
colonist, C. cyanogutattum. The associated fish fauna of the basin has been 
described by Minckley (1969, 1978). 

Etymology.—The species is named after Dr. W. L. Minckley of Arizona State 
University who has investigated the biota of Cuatro Ciénegas for many years. 


Acknowledgments 


We sincerely appreciate the guidance of C. D. N. Barel in our morphological 
analyses. Invaluable assistance was provided by David C. Smith. Our thanks to 
Hugh DeWitt for reading the manuscript; E. C. Theriot and W. N. Watkins 
prepared Figures | and 2, respectively. We gratefully acknowledge the cooper- 
ation of Carlos Tovar Flores, Of. Coordinadora de Investigaciones Pesqueras 
Externas, Direccion General del Instituto Nacional de la Pesca, for permitting 
our field work in Cuatro Ciénegas. This research was supported by NSF grant 
DEB 78-24074 (IK), the Migratory Fish Research Institute, Faculty Research 
Fund, University of Maine, and Earthwatch, The Center for Field Research, 
Belmont, Massachusetts. 


Literature Cited 


Baerends, G. P., and J. M. Baerends-Van Roon. 1950. An introduction to the study of the ethology 
of cichlid fishes.—Behaviour, Supplement number 1:1-243. 


268 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Barel, C. D. N., M. J. P. van Oijen, F. Witte, and E. L. M. Witte-Maas. 1977. An introduction to 
the taxonomy and morphology of the haplochromine Cichlidae from Lake Victoria.—Nether- 
lands Journal of Zoology 27:333-389. 

Deacon, J. E., G. Kobetich, J. D. Williams, S. Contreras, and others. 1979. Fishes of North 
America—endangered, threatened, or of special concern: 1979.—Fisheries 4:29—44. 

Graves, J. E., and R. H. Rosenblatt. 1980. Genetic relationships of the color morphs of the serranid 
fish Hypoplectrus unicolor.—Evolution 34:240-245. 

Hoogerhoud, R. J. C., and C. D. N. Barel. 1978. Integrated morphological adaptations in piscivo- 
rous and mollusc-crushing Haplochromis species.—Proceedings of the Zodiac Symposium on 
Adaptation (Pudoc Wageningen, the Netherlands):52—56. 

Hubbs, C. L., and R. R. Miller. 1965. Studies of cyprinodont fishes. XXII. Variation in Lucania 
parva, its establishment in western United States, and description of a new species from an 
interior basin in Coahuila, Mexico.—Miscellaneous Publications of the Museum of Zoology, 
University of Michigan 127:1-111. 

Hutchinson, G. E. 1978. An introduction to population ecology.—Yale University Press, New 
Haven, Connecticut, 260 pp. 

Jimenez, G. F., G. Guajardo M., and C. H. Briseno. 1981. Trematodos de peces dulceacuicolas de 
Coahuila, Mexico. I. Quadripaludis luistoddi gen. et sp. nov. (Trematoda: Hemiuridae) par- 
asitos de ciclidos endemicos de Cuatro Ciénegas.—Southwestern Naturalist 26:409-413. 

Kornfield, I. L. 1978. Evidence for rapid speciation in African cichlid fishes.—Experientia 34:335— 
336. 

—. 1981. Biological status of the cichlid fishes of Cuatro Ciénegas.—Proceedings of the Desert 

Fishes Council 12:96—97. 

,and R. K. Koehn. 1975. Genetic variation and speciation in New World cichlids.—Evolution 

94:427-437. 

, D. C. Smith, P. S. Gagnon, and J. N. Taylor. 1982. The cichlid fish of Cuatro Ciénegas, 

Mexico: direct evidence of conspecificity among distinct trophic morphs.—Evolution 36:658— 

664. 

LaBounty, J. F. 1974. Materials for the revision of cichlids from northern Mexico and southern 
Texas, U.S.A.—Unpublished Ph.D. dissertation, Arizona State University, Tempe, Arizona, 
121 pp. 

Liem, K. F. 1979. Modulatory multiplicity in the feeding mechanism in cichlid fishes, as exemplified 
by the invertebrate pickers of Lake Tanganyika.—Journal of Zoology 189:93-125. 

Meek, S. E. 1904. The fresh-water fishes of Mexico north of the Isthmus of Tehuantepec.—Field 
Columbia Museum Publication 93 (Zoology Series, 5):1—252. 

Miller, R. R. 1968. Two new species of the genus Cyprinodon from the Cuatro Ciénegas basin, 
Coahuila, Mexico.—Occasional Papers of the Museum of Zoology, University of Michigan 
659: 1-15. 

Minckley, W. L. 1969. Environments of the bolson of Cuatro Ciénegas, Coahuila, Mexico, with 
special reference to the aquatic biotaa—Texas Western Press, University of Texas, El Paso, 
Texas, Science Series 2:1-63. 

——. 1978. Endemic fishes of the Cuatro Ciénegas Basin, Northern Coahuila, Mexico. Pp. 383- 
404 In: R. H. Wauer and D. Riskind, eds., Trans. Symp. Biol. Resources of the Chihuahuan 
Desert Region, U.S. and Mexico.—Transactions and Proceedings of the U.S. National Parks 
Service. Ser. 3. York. 

Regan, C. T. 1906-08. Pisces. In: Biologia Centrali-Americana 8: 1—203. 

Sage, R. D., and R. K. Selander. 1975. Trophic radiation through polymorphism in cichlid fishes.— 
Proceedings of the National Academy of Sciences 72:4669-4673. 

Smith, D. C. 1982. Trophic ecology of the cichlid morphs of Cuatro Ciénegas, Mexico.—Unpub- 
lished M.S. thesis, University of Maine, Orono, Maine, 45 pp. 

Taylor, D. W. 1966. A remarkable snail fauna from Coahuila, Mexico.—The Veliger 9:152—228. 

, and W. L. Minckley. 1966. New World for biologists.—Pacific Discovery 19:18—22. 

Taylor, J. N., and R. R. Miller. 1980. Two new cichlid fishes, genus Cichlasoma, from Chiapas, 

Mexico.—Occasional Papers of the Museum of Zoology, University of Michigan 693: 1-16. 

, and (In press). Cichlid fishes (genus Cichlasoma) of the Rio Panuco basin, eastern 

Mexico, with description of a new species.—Occasional Papers of the Museum of Natural 

History, University of Kansas. 


VOLUME 96, NUMBER 2 269 


Thompson, K. W. 1979. Cytotaxonomy of 41 species of neotropical Cichlidae.—Copeia 1979:679- 


691. 
Turner, B. J., and D. J. Grosse. 1980. Trophic differentiation in J/yodon, a genus of stream-dwelling 


goodeid fishes: speciation versus ecological polymorphism.—Evolution 34:259-270. 
Vermeij, G. J., and A. P. Covich. 1978. Coevolution of freshwater gastropods and their predators.— 
American Naturalist 112:833-843. 


(IK) Department of Zoology and Migratory Fish Research Institute, University 
of Maine, Orono, Maine 04469; (JNT) Department of Biological Sciences, Florida 
Atlantic University, Boca Raton, Florida 33432. 


PROC. BIOL. SOC. WASH. 
96(2), 1983, pp. 270-272 


CLARIFICATION OF THE NAMES RANA MYSTACEA 
SPIX, 1824, LEPTODACTYLUS AMAZONICUS 
HEYER, 1978 AND A DESCRIPTION OF A 
NEW SPECIES, LEPTODACTYLUS SPIXI 
(AMPHIBIA: LEPTODACTYLIDAE) 


W. Ronald Heyer 


Abstract.—Méhely in 1904 proposed a lectotype designation for Rana mysta- 
cea Spix, 1824, that differs from Heyer’s lectotype designation of 1978. Lepto- 
dactylus amazonicus Heyer, 1978, is a synonym of Rana mystacea Spix, 1824, 
and a new species is proposed to solve the remaining consequent nomenclatural 
problem. 


Dr. Marinus Hoogmoed brought to my attention a paper by Méhely (1904) 
which I had overlooked in my review of the Leptodactylus fuscus species group 
(Heyer 1978). Méhely (1904: 219) designated a lectotype for Rana mystacea 
Spix, 1824, which differs from and obviously predates my designation for the 
species (Heyer 1978:30). To summarize briefly, Spix based his new species, mys- 
tacea, on two Brazilian specimens (since destroyed), a male from “‘Bahia”’ (Spix 
1824, plate 3, figure 3) and a female from ‘‘Solimoens’’ (=Rio Solimoes) (Spix 
1824, plate 3, figure 1). Méhely (1904:219) considered the Bahia specimen to 
belong to the species Rana typhonia Daudin [=Leptodactylus fuscus (Schneider, 
1799)] and chose the Rio Solimoes specimen as represented in figure 1, plate 3 
of Spix to be the name-bearer of mystacea. I considered the Bahia and Solimoes 
specimens to represent two distinct members of the mystaceus complex and chose 
the figure of the Bahia specimen as the name bearer of mystacea and described 
a new species, amazonicus, for the species represented by the figured specimen 
from the Rio Solimoes. As Méhely’s action takes precedence, the consequences 
are: (1) Leptodactylus amazonicus Heyer, 1978, becomes a synonym of Rana 
mystacea Spix, 1824; (2) the species of the mystaceus complex from the northern 
Atlantic forests of Brazil lacks a name, as no other name has been proposed for 
the species involved. Consequently, the following name is proposed for the species 
referred to as Leptodactylus mystaceus in the 1978 paper. 


Leptodactylus spixi, new species 


Holotype. —USNM 96409, an adult male from Brazil: Rio de Janeiro; Saco de 
Sao Francisco, Niteroi. Collector unknown, collected on 13 Oct 1923. 

Paratopotypes.—USNM 96407-8, 96410-11. 

Referred specimens.—Other specimens as indicated under distribution section 
for Leptodactylus mystaceus in Heyer, 1978 (p. 65). 

Diagnosis.—Most individual spixi have a combination of a distinct light stripe 
on the posterior surface of the thigh and distinct white tubercles on the surfaces 
of the posterior tarsus and sole of foot; these states are shared with albilabris 


VOLUME 96, NUMBER 2 271 


(Gunther), 1859; elenae Heyer, 1978; fragilis (Brocchi), 1877; and latinasus (Es- 
pada), 1875. Leptodactylus spixi has distinct dorsolateral folds (at least indicated 
by color pattern), fragilis and latinasus lack distinct dorsolateral folds. Lepto- 
dactylus spixi has white tubercles on the dorsal surface of the tibia; the tibia is 
smooth in elenae. Leptodactylus spixi is found in east coastal Brazil, albilabris 
occurs in the West Indies. 

Some individuals of L. spixi lack the white tubercles on the tarsus and sole of 
foot (light thigh stripe present). These states are shared with at least some indi- 
viduals of furnarius Sazima & Bokermann, 1978 (=/aurae Heyer, 1978), fuscus 
(Schneider), 1799, geminus Barrio, 1973, gracilis (Duméril & Bibron), 1841, lon- 
girostris Boulenger, 1882, notoaktites Heyer, 1978, and poecilochilus (Cope), 
1862. The tubercles on the dorsal surface of the tibia distinguish L. spixi from all 
these species. 

Description of holotype.—Snout shape subelliptical from above, rounded with 
protruding ridge from side; canthus rostralis indistinct; loreal slightly concave in 
cross section; tympanum large, distinct, diameter about 34 eye diameter; well 
developed vocal slits present; slight external lateral vocal fold present, not de- 
veloped into well developed sac; vomerine teeth in two long angulate, arched 
series almost in medial contact posterior to choanae; first finger much longer than 
second; fingers I and IV free, fingers II and III with slight lateral ridges; thumb 
lacking asperities; dorsum smooth; a pair of well developed dorsolateral folds 
from behind eye to hind leg; distinct supratympanic fold, from eye to shoulder; 
belly smooth, ventral disk distinct; toes free, tips not expanded; subarticular 
tubercles moderately developed; metatarsal ridge barely indicated; tarsal fold 
extending about % length tarsus; upper tibia scattered with white-tipped tuber- 
cles; posterior surface of tarsus and sole of foot with many white-tipped tubercles. 

SVL 43.0 mm, head length 19.8 mm, head width 14.5 mm, interorbital distance 
3.2 mm, eye-nostril distance 4.2 mm, femur 16.4 mm, tibia 21.5 mm, foot 
24.7 mm. 

Dorsum with irregular darker brown markings on lighter brown background 
including irregular interorbital triangle with apex directed posteriorly and chevron 
in suprascapular region; dorsolateral folds barely highlighted by darker brown 
color; dark canthal stripe from nostril to tympanum, interrupted by eye; distinct 
light stripe from tip of snout, running below dark canthal stripe, under eye and 
tympanum to angle of jaw; upper lip with slightly darker shading below light 
stripe; limbs barred above; venter immaculate; posterior surface of thigh almost 
uniform tan dorsally, bordered below by distinct light stripe highlighted by dark 
brown above and below. 

Etymology.—Named for J. B. Spix, who was not to blame for the confusion 
that his figures have caused subsequent generations of herpetologists. 

Distribution:—To my knowledge, no additional specimens have been docu- 
mented from east coastal Brazil than those listed previously (Heyer 1978:65, and 
fig. 61). 


Acknowledgments 


Dr. Marinus Hoogmoed (Rijksmuseum van Natuurlijke Historie, Leiden) not 
only pointed out the Méhely paper I had overlooked, but analyzed the nomen- 


272 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


clatural situation in consultation with his colleague, Dr. Holthius, and suggested 
that I should propose a new species to solve the situation. 

Ronald I. Crombie and George R. Zug (Smithsonian Institution, USNM) kindly 
reviewed this paper. 


Literature Cited 


Heyer, W. R. 1978. Systematics of the fuscus group of the frog genus Leptodactylus (Amphibia, 
Leptodactylidae).—Science Bulletin of the Natural History Museum of Los Angeles County 
29: 1-85. 

Méhely, L. 1904. Investigations on Paraguayan batrachians.—Annales Historico-Naturales Musei 
Nationalis Hungarici 2:207—232 + Plate XIII. 

Spix, J. B. 1824. Animalia nova sive species novae Testudineum et Ranarum, quas in itenere per 
Brasiliam, annis 1817-1820. Monaco. 53 pp. + 22 plates. 


Department of Vertebrate Zoology, Smithsonian Institution, Washington, D.C. 
20560. 


PROC. BIOL. SOC. WASH. 
96(2), 1983, pp. 273-275 


A NEW BATHYAL SPECIES OF CORALLIODRILUS 
(OLIGOCHAETA: TUBIFICIDAE) FROM 
THE SOUTHEAST ATLANTIC 


Christer Erséus 


Abstract.—Coralliodrilus longiductus, n. sp. (subfamily Phallodrilinae) is de- 
scribed from the continental slope west of Angola. It is the first species of Cor- 
alliodrilus from bathyal waters, and it is distinguished from its congeners by its 
slender, bipartite, and heavily muscular atria. 


The genus Coralliodrilus was established by Erséus (1979) for C. leviatriatus 
from coral reef sands at Bermuda. Subsequently, seven additional species, four 
from the Great Barrier Reef (Erséus 1981), and three from Italy (Erséus, in press), 
have been described. 

While examining a collection of oligochaetes from the “‘Atlantis II-42”’ cruise 
undertaken by the Woods Hole Oceanographic Institution in Walvis Bay (SE 
Atlantic), a new species closely related to the shallow-water species of Coral- 
liodrilus was encountered. This note provides the description of this species. 

The specimens were all stained in paracarmine and mounted whole in Canada 
balsam. The type-material has been deposited in the National Museum of Natural 
History (USNM), Smithsonian Institution, Washington, D.C. 


Coralliodrilus longiductus, new species 
Fig. 1 


Holotype.—USNM 74635 

Type-locality.—Continental slope west of Luanda, Angola, 09°05’S, 12°17’E to 
08°56’'S, 12°15’E, 1427-1643 m (AII-42 Sta. No. 202). 

Paratypes.—USNM 74636-74639, 4 whole-mounted specimens from the type- 
locality. 

Other material examined (author’s collection).—Three whole-mounted speci- 
mens from the type-locality. 

Description.—Length (2 complete, fixed specimens) 2.8-3.1 mm, 33-34 seg- 
ments. Diameter at clitellum (slightly compressed worms), 0.14—-0.18 mm. Pro- 
stomium small. Clitellum extending over 12X—XII. Somatic setae (Fig. 1A) bifid, 
with upper tooth thinner and shorter than lower. Bifids slender, 32-44 um long, 
about | «wm thick, 3 per bundle anteriorly, 2-3(4) per bundle in post-clitellar 
segments. Ventral setae of XI modified into penial bundles (Fig. 1B, ps), each of 
which contains 5—7 (generally 6) straight setae, 25-33 wm long, about 1.5 wm 
thick. Ectal tips of penial setae very narrow, single-pointed and curved, protrud- 
ing into male invaginations. Male pores as paired invaginations (Fig. 1B, mi), 
located in line with ventral somatic setae, in posterior part of XI. Spermathecal 
pores paired in lateral lines, in anterior part of X. 

Pharyngeal glands extending into VI. Male genitalia (all structures paired) (Fig. 
1B): whole vas deferens not observed in available material, but appears shorter 


274 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 1. Coralliodrilus longiductus: A, Free-hand drawing of somatic seta; B, Lateral view of 
spermatheca and male genitalia in segments X—XI. aa, atrial ampulla; ad, atrial duct; mi, male invagi- 
nation; pp, pseudopenis; ps, penial setae; s, spermatheca; sf, sperm funnel; vd, vas deferens. 


than atrium and is, ectally at least, muscular; vs entering apical, ental end of 
atrium; atrial ampulla, 90-115 wm long, 22-32 wm wide, with 3-4 wm thick outer 
lining of longitudinal muscles, and ciliated, granulated inner epithelium; atrial 
duct, up to 80 wm long, about 10 wm wide, muscular, terminating in small, mus- 
cular, protrusible pseudopenis in lateral wall of male invagination; prostate glands 
absent. Spermathecae (Fig. 1B, s) with ducts, 40-55 wm long, 13-25 wm wide, 
and very thin-walled ampullae, 50-85 wm long, 25-45 wm wide; sperm in random 
masses. 

Remarks.—This new species shares the feature of bipartite atria with the Aus- 
tralian C. atriobifidus and C. oviatriatus (both described by Erséus 1981), and 
one of the new species from Italy (Erséus, in press). However, C. longiductus, 
n. sp. is distinguished from the other three by (1) its heavily muscular atria (atrial 
musculature thin in atriobifidus and oviatriatus, distinct but of varying thickness 
in the Italian species), and (2) its slender spermathecal ducts (these ducts are 
elaborate and longer than the spermathecal ampullae in C. atriobifidus, short and 
inconspicuous in C. oviatriatus and the Italian species). 


Acknowledgments 
I am indebted to Drs. H. L. Sanders, F. J. Grassle, and G. R. Sampson (WHOD), 
for placing the material at my disposal. 
Literature Cited 


Erséus, C. 1979. Coralliodrilus leviatriatus gen. et sp.n., a marine tubificid (Oligochaeta) from 
Bermuda.—Sarsia 64:179-182. 


VOLUME 96, NUMBER 2 275 


1981. Taxonomic studies of Phallodrilinae (Oligochaeta, Tubificidae) from the Great Barrier 
Reef and the Comoro Islands with descriptions of ten new species and one new genus.— 
Zoologica Scripta 10:15-31. 


. (In press). Three new species of the marine genus Coralliodrilus (Oligochaeta, Tubificidae) 
from Italy.—Bolletino di Zoologia. 


Swedish Museum of Natural History, Stockholm, and Department of Zoology, 


University of Gothenburg; postal address: Department of Zoology, Box 25059, 
S-400 31 Gothenburg, Sweden. 


PROC. BIOL. SOC. WASH. 
96(2), 1983, pp. 276-290 


NEW SPECIES OF FABRICIOLA AND FABRICIA 
(POLYCHAETA: SABELLIDAE) FROM BELIZE 


Kirk Fitzhugh 


Abstract.—Two new species of sabellid polychaetes, Fabriciola trilobata and 
Fabricia infratorquata, are described from Belize. Methyl green staining patterns 
of epithelial mucous gland cells are also described. Setal and collar segment 
characteristics are discussed. 


An analysis of benthic samples from Twin Cays, Belize, revealed the presence 
of two numerically dominant undescribed species belonging to the genera Fabri- 
ciola Friedrich, 1939, and Fabricia Blainville, 1828. These species are described 
in the present paper. I also give the first report of methyl green staining and its 
variability in these genera. Staining in sabellids has been reported by Banse (1970, 
1972, 1979) for a few genera, but not in Fabriciola or Fabricia. In his work on 
species of Euchone, Banse (1970) suggested the importance of this technique in 
providing additional diagnostic characters. 

The present descriptions and references to other species follow the pattern 
established by Banse (1956), according to whom the development of the collar 
dorsally and ventrally is the primary character distinguishing the two genera. 
This pattern was continued by Banse (1957) and Hartmann-Schroeder (1971). In 
the species key provided by Friedrich (1939), shaft length of abdominal uncini 
was used as a main diagnostic feature. Diagnoses given by Day (1967) included 
collar characteristics plus uncini shaft length and condition of palps (filamentous 
or not); the two latter characters appeared to be given greater weight. Fauchald 
(1977) restated Day’s diagnoses. In Fauchald’s key, genera were separated on 
the basis of possession of long- or short-handled abdominal uncini. The two 
genera cannot be separated based on shaft length since both long and short forms 
occur in both genera. Collar development as put forth by Banse (1956) is thus 
suggested as a more useful and consistent character by which to distinguish the 
genera Fabriciola and Fabricia. 


Fabriciola trilobata, new species 
Figs. 1-2, 3a—c; Table 1 


Material examined.—West Bay, Twin Cays, Belize; 30 cm depth; mat of Cau- 
lerpa verticillata on rootmat of Rhizophora mangle, some organic debris and 
fragments of Halimeda; 7 Apr 1982 (Array F202), 9 Apr 1982 (Array F203), 11 
Apr 1982 (Array F204); coll. K. Fauchald. Holotype: F204 A-4 (USNM 74679). 
Paratypes: F204 A-1, 4 specimens (USNM 74680); F204 A-2, 6 specimens (USNM 
74681); F204 A-3, 9 specimens (USNM 74682); F204 A-4, 13 specimens (UNSM 
74683); F204 A-5, 5 specimens (UNSM 74684); F204 B-1, 11 specimens (USNM 
74685); F204 B-2, 16 specimens (USNM 74686); F204 B-3, 7 specimens (USNM 
74687); F204 B-4, 8 specimens (USNM 74688); F204 B-5, 6 specimens (USNM 
74689); F204 C-1, 1 specimen (USNM 74690); F204 C-2, 11 specimens (USNM 


VOLUME 96, NUMBER 2 ZT 


74691); F204 C-3, 6 specimens (USNM 74692); F204 C-4, 2 specimens (USNM 
74693); F204 C-5, 7 specimens (USNM 74694); F204 D-1, 2 specimens (USNM 
74695); F204 D-2, 1 specimen (USNM 74696); F204 D-3, 9 specimens (USNM 
74697); F204 D-4, 10 specimens (USNM 74698); F204 D-5, 8 specimens (USNM 
74699). Additional material: F202 A-1, 4 specimens (USNM 74700); F202 A-3, 1 
specimen (USNM 74701); F202 A-4, 4 specimens (USNM 74702); F202 A-5, 4 
specimens (USNM 74703); F202 B-1, 4 specimens (USNM 74704); F202 B-2, 1 
specimen (USNM 74705); F202 B-4, 8 specimens (USNM 74706); F202 C-1, 3 
specimens (USNM 74707); F202 C-2, 6 specimens (USNM 74708); F202 C-3, 3 
specimens (USNM 74709); F202 C-4, 1 specimen (USNM 74710); F202 C-5, 4 
specimens (USNM 74711); F202 D-1, 1 specimen (USNM 74712); F202 D-2, 5 
specimens (USNM 74713); F202 D-3, 4 specimens (USNM 74714); F202 D-5, 1 
specimen (USNM 74715); F203 A-1, 2 specimens (USNM 74716); F203 A-2, 2 
specimens (USNM 74717); F203 A-3, 1 specimen (USNM 74718); F203 A-4, 4 
specimens (USNM 74719); F203 A-5, 4 specimens (USNM 74720); F203 B-1, 2 
specimens (USNM 74721); F203 B-2, | specimen (USNM 74722); F203 B-3, 4 
specimens (USNM 74723); F203 B-4, 1 specimen (USNM 74724); F203 B-5, 2 
specimens (USNM 74725); F203 C-1, 1 specimen (USNM 74726): F203 C-2, 2 
specimens (USNM 74727); F203 C-3, 1 specimen (USNM 74728); F203 C-4, 6 
specimens (USNM 74729); F203 C-5, 2 specimens (USNM 74730); F203 D-1, 2 
specimens (USNM 74731); F203 D-2, 6 specimens (USNM 74732); F203 D-3, 2 
specimens (USNM 74733); F203 D-4, 6 specimens (USNM 74734); F203 D-5, 2 
specimens (USNM 74735). 

Diagnosis.—Fabriciola with filiform palps of variable length. Peristomium re- 
stricted to dorsal half of anterior end; visible ventrally and laterally, fused dor- 
solaterally to collar segment; anterior ventral margin expanded into shelf-like 
process. Ventral collar smooth, fused dorsolaterally to collar segment. Dorsal 
collar higher, divided by middorsal longitudinal groove. Anterior groove margin 
extended as triangular lobe, overlapped posteriorly by pair of collar lobes. Tho- 
racic uncini with main fang surmounted by large offset tooth and 2 smaller teeth, 
crested by 4 rows of smaller teeth. Short-handled abdominal uncini with rows of 
teeth increasing in number with successive setigers: 7, 7-8 and 9 rows of teeth 
in setigers 9, 10 and 11, respectively. Abdominal neurosetae include 1—2 superior 
minute limbate setae and 2-3 inferior long, narrow limbate setae. 

Description.—The holotype is a complete specimen with 9 thoracic and 3 ab- 
dominal setigers. Length 4.04 mm (0.78 mm comprising branchial crown) and 
width 0.19 mm. 

Attached to the peristomium are a pair of semicircular branchial lobes, each 
with a large branchial heart located dorsally (Fig. la). Three radioles are attached 
to each branchial lobe; they are unbranched and rounded externally. Usually 7 
pairs of ciliated pinnules extend from the inner side of each radiole. The proximal 
pair are longest, with more distal pinnules becoming shorter, such that all extend 
to about the same height; nearly 34 the radiole length. Distal ends of the radioles 
are drawn out as very fine filaments. In the holotype the filamentous end con- 
tributes about 4 of the radiole length, but in paratypes examined the end may 
be as much as '% the radiole length. A pair of filiform palps are present, each 
originating from the inner side of each branchial lobe, adjacent to the middle 
radiole. Palps are distally blunt, and, in the holotype are about 34 the radiole 


278 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Table 1.—Comparison of selected species of Fabriciola. 


Species Peristomium Collar Thoracic notosetae 
trilobata Visible laterally and ven- Smooth, even ventrally; Setiger 1-8: 4—5 long lim- 
trally; limited to dorsal higher dorsally, middor- bates. Setiger 3-8: 1-2 
half of anterior end. sal margin trilobate. spatulates. 
capensis Visible dorsally, ventrally Smooth, even ventrally; Long bilimbate setae. 
and laterally. dorsally same height as 


ventrum, middorsal 
margin trilobate. 


length. In paratypes, palp length is variable, ranging from '4 to %4 the radiole 
length. Internally, the length of the palps is almost completely occupied by a 
convoluted vessel, representing an extension of either the circulatory system or 
coelom. 

The peristomium is visible ventrally and laterally (Fig. 1b, c), limited to the 
dorsal half of the anterior end. The ventral anterior margin is complete and slightly 
expanded, forming a rounded shelf-like process. Dorsolaterally, the peristomium 
fuses with the collar segment. 

A distinct collar is present. Ventrally it is entire, even, and with rounded edges. 
Dorsolaterally, it fuses with the collar segment. Dorsally the collar extends slight- 
ly further anteriorly than it does ventrally. A middorsal longitudinal groove splits 
the anterior 4 of the collar segment into right and left halves. A small triangular 
lobe extends from the anterior end of the groove and Is partly overlapped pos- 
teriorly by a pair of lateral lobes, each of which is slightly larger than the median 
lobe. 

The collar segment is basically cylindrical, becoming only slightly wider pos- 
teriorly. The intersegmental groove between the collar segment and setiger | is 
only distinct ventrally. Just anterior to the groove is a partial annulation, complete 
ventrally and disappearing laterally. It is assumed this does not denote an addi- 
tional achaetous segment. 

Thoracic setigers are cylindrical with distinct segmentation. Setiger 1 is short- 
est, about 12 the length of the collar segment, and slightly wider than long. Each 
subsequent thoracic setiger is longer than the previous. Setiger 2 is as long as 
wide, setiger 5 is about twice as long as wide, and setiger 8 about 2.5 times as 
long as wide. 

The abdominal setigers (9-11) become progressively shorter (Fig. 2a). Setigers 
9-10 are cylindrical, 11 is dorsoventrally flattened. Setiger 9 is %4 as long as setiger 
8 and 1.5 times longer than wide. Setiger 10 is as long as wide and setiger 11 is 
wider than long. Setigers 9-10 are the same width as thoracic segments, with 
setiger 11 abruptly narrower. Segmentation is distinct. Swollen, pad-like glandular 
areas are present just posterior to notopodial tori of setiger 9-10 and at anterior 
lateral margins of setiger 11. 

Thoracic noto- and neuropodial tori are represented only by a slight swelling. 
Notopodia of setiger | are situated dorso-laterally; neuropodia are absent. No- 


VOLUME 96, NUMBER 2 279 


Table 1.—Continued. 


Thoracic neurosetae Abdominal notosetae Abdominal neurosetae 
5-7 uncini. Main fang + 1 Short-handled uncini. Setiger |—2 superior minute, narrow 
large and 2 smaller teeth + 9: 25 uncini; 7 rows of limbates; 2—3 inferior long, 
4 arcs of smaller teeth. teeth. Setiger 10: 28 uncini; narrow limbates. 


7-8 rows of teeth. Setiger 
11: 18 uncini; 9 rows of 


teeth. 
Uncini with main fang + 1-2 Long-handled uncini with 18 Slender capillaries. 
large teeth + 1 arc of 12-14 rows of teeth. 


smaller teeth. 


topodia of subsequent thoracic setigers are lateral. Neuropodia are ventral to and 
slightly posterior to notopodia. Notosetae in setiger 1 include 4—S long superior 
and 1 shorter inferior limbate setae. Superior notosetae in setigers 2-8 are long 
limbates similar to those of setiger 1, numbering 4—5 (Fig. 2d). Setigers 3-8 also 
possess 1-2 inferior short spatulate setae (Fig. 2b). Thoracic neurosetae are gently 
curved acicular uncini; 5—7 per fascicle, in a single vertical row (Fig. 2g). In lateral 
view, the distal ends of uncini have a large main fang surmounted obliquely by 
a large tooth followed by 4 smaller teeth. In frontal view, the large tooth above 
the main fang is clearly offset and accompanied by 2 smaller teeth (Fig. 2h). Teeth 
above this first row form a series of concentric arcs. Proximal to the distal end 
is a Slight swelling of the shaft, which tapers proximally to a rounded end. 

Abdominal notosetae occur on distinct lateral tori located on the posterior 4 
of the setiger, and are the same length as thoracic neuropodia. The number of 
uncini varies in that setiger 9 has 25, setiger 10 has 28 and setiger 11 has 18 uncini. 
Uncini are short-handled with a slight constriction below the proximal-most tooth 
row, followed by a slight swelling and a truncate base (Fig. 21). Uncini of setiger 
9 have 7 rows of teeth; the number of teeth per row from proximal to distal as 
follows: 1+2+3+3+4+6+5. Setiger 10 uncini have 7-8 tooth rows, those with 8 
rows having the formula: 1+2+3+3+4+4+4+4. Setiger 11 uncini possess 9 tooth 
rows with the formula: 1+2+4+3+4+3+4+4+4. Abdominal neurosetae originate 
just ventral to notopodial tori. No neuropodial tori could be discerned. Neuro- 
setae are of 2 types: 1-2 very minute superior limbate setae with a narrow lim- 
bation (Fig. 2c) and 2-3 inferior limbate setae with very long, slender shafts and 
a very narrow limbation extending about %4 the length of the shaft (Fig. 2e-f). 
Neurosetae are directed dorsally. 

The pygidium is roughly triangular and dorsoventrally flattened (Fig. 2a). An- 
teriorly it is the same width as the adjacent segment, narrowing and becoming 
rounded posteriorly. A V-shaped glandular zone extends along the outer margin, 
the arms of the V becoming wider posteriorly. The anus is a depressed, mid- 
ventral longitudinal slit. 

A pair of dark crescentic-shaped eyes (dorsal view) are in the collar segment 
on either side of the middorsal groove. A pair of smaller, circular eyes are located 
laterally in the posterior 4 of the pygidium. No otocysts could be found. 

The holotype (in alcohol) is opaque and cream colored. The palps of some 


280 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


a 


0.2 mm a-c b C 


Fig. 1. Fabriciola trilobata (holotype, USNM 74679): a—c, Dorsal, ventral and lateral views of 
anterior end. 


individuals examined were dull orange or yellow, sometimes with a corresponding 
lighter coloration in the radioles. Some individuals contained dark brown pig- 
mentation extending dorsally on the branchial lobes, ventral peristomial margin, 
and along the anterior margin of the ventral collar. The middorsal groove, adja- 
cent posterior region and lateral lobes showed similar pigmentation. 

Tubes are thick, about 2—2.5 times as wide as the individual, constructed with 
loosely bound plant and detrital material. The central space within the tube is 
very narrow, such that nearly the entire length of the worm is in contact with the 
tube wall. 


VOLUME 96, NUMBER 2 281 


0.016 mm 


Fig. 2. Fabriciola trilobata (a from holotype; b-i from paratype, USNM 74683): a, Posterior end, 
dorsal view; b, Thoracic spatulate seta from setiger 3; c, Superior abdominal neuroseta from setiger 
10; d, Thoracic notoseta from setiger 3; e-f, Proximal and distal portions of inferior abdominal neu- 
roseta from setiger 10; g—h, Lateral view of thoracic neuropodial uncinus and frontal view of tooth 
arrangement, both from setiger 5; 1, Abdominal notopodial uncinus from setiger 9. 


The holotype and some paratypes were stained with a dark solution of methyl 
green in ethanol. Specimens were placed in the solution for 30 minutes, allowed 
to destain in ethanol for 10 minutes and observed. Uptake of stain differed con- 
siderably from holotype and paratypes; thus both patterns will be described. 

In the holotype the collar segment only stained ventrally, extending from just 
posterior to the collar margin to the posterior segment margin, becoming darker 


282 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


posteriorly (Fig. 3a). Setiger 1 stained darkest ventrally along the anterior margin, 
extending posteriorly as a triangular pattern to the near the ventral midpoint. 
Remainder of the ventrum was more lightly stained, fading laterally and dorsally. 
Notopodial tori did not stain. On setiger 2 a broad midventral strip was darkest, 
with less staining to either side, fading laterally and dorsally. Noto- and neuro- 
podial tori of this and following thoracic and abdominal setigers did not stain. 
Setiger 3 stained evenly ventrally, with lateral and dorsal staining as in previous 
setigers. The ventral anterior half of setiger 4 was well stained, becoming lighter 
posteriorly, laterally and dorsally. Setigers 5—7 had fewer, but larger, stain-ac- 
cepting cells ventrally and laterally, thus imparting a light green hue. The anterior 
margin of setiger 8 was stained as in adjacent setigers, becoming darker poste- 
riorly. Setigers 9-10 were darker and evenly stained on both sides of the segments 
with large staining cells. Setiger 11 was similar to 9-10 but darker. Glandular 
areas on abdominal setigers stained darker than the rest of the corresponding 
segment. The pygidium stained very deeply, especially the glandular area. The 
fecal groove was non-staining, extending from the anterior margin of the anus 
along the ventral midline of setigers 11-10. On setiger 9 it extends diagonally to 
the left to near the dorsal midline of the anterior margin, continuing anteriorly to 
the collar segment. 

In paratypes examined, the anterior portion of the collar segment accepted 
stain only in a well defined lateral area just posterior to the eyes, extending to 
near the dorsal midline posteriorly (Fig. 3b—c). Much of this area was character- 
ized by large, dark staining cells. The posterior region of the collar segment was 
uniformly stained ventrally and ventrolaterally. Setiger 1 showed a dense staining 
region along the midventral anterior margin, with the remainder of the ventrum 
lightly stained. Laterally, a green hue was present in addition to some large stain- 
accepting cells. The anterior and posterior midventral margins of setiger 2 were 
densely stained, in addition to large staining cells interspersed between the two 
areas. The remaining ventral and lateral areas were lightly stained. The ventral 
anterior margin of setiger 3 was darkly stained, becoming lighter posteriorly with 
some large staining cells. Lateral staining was similar to anterior setigers. The 
remaining segments and pygidium stained similar to the holotype. 

Remarks.—Fabriciola trilobata shows some similarity to F. capensis (Monro 
1937:366; see also Day 1955:447). Table 1 compares the two species. Both exhibit 
a trilobate middorsal collar margin formed by the presence of a middorsal groove. 
The body of both species is elongate, but F. capensis possesses much longer 
segments. The two species differ in that in F. trilobata the dorsal side of the 
collar is higher than the ventral, while in F. capensis it is even; also, all thoracic 
notosetae in the latter species are bilimbate and no spatulate setae have been 
reported. Abdominal uncini of the two species differ in number of tooth rows (7— 
9 in F. trilobata and 18 in F. capensis). 

In addition to the above characters, the tooth arrangement above the main fang 
of thoracic uncini, increasing number of tooth rows in uncini of successive ab- 
dominal setigers, and the presence of minute superior limbate setae in abdominal 
neuropodia have not been described in any other Fabriciola species. Banse (1957) 
reported the presence of minute ‘‘needle-like’’ setae in thoracic notopodia of 
Oriopsis rivularis (Annenkova 1929), and Banse (1959) noted a similar setal type 


VOLUME 96, NUMBER 2 283 


Fig. 3. Fabriciola trilobata: a, Staining pattern of anterior end of holotype, ventral view; b—c, 
Staining pattern of anterior end of paratype (USNM 74685), ventral and lateral views. Fabricia infra- 
torquata (all from paratypes, USNM 74658): d—e, Superior and inferior thoracic notosetae from setiger 
6; f, Abdominal neuroseta from setiger 9; g—h, Lateral view of thoracic neuropodial uncinus and 
frontal view of tooth arrangement, both from setiger 6; i, Thoracic spatulate seta from setiger 3; j, 
Abdominal notopodial uncinus from setiger 9. 


284 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Table 2.—Comparison of selected species of Fabricia. 


Species Palps Collar Thoracic notosetae 

infratorquata Absent Indistinct dorsally, split by Setiger 1-8: 3-4 long limbates. 
middorsal groove; large Setiger 2,6—-8: 1 short limbate. 
triangular lobe ventrally. Setiger 3-5: 1-2 spatulates. 

bansei Absent Indistinct dorsally, large Setiger 1-8: 4—5 limbates. Setiger 
triangular lobe ventrally. 2-8: 2—3 spatulates. 

brunnea Absent Pair semicircular lobes dor- _Setiger 1: pointed setae. Setiger 
sally; elongate lobe ven- 2-8: 3 limbates, ! spatulate. 
trally. 

gerdi Absent Indistinct dorsally, large 4-6 long limbates, 1—2 short lim- 
rounded lobe ventrally. bates. 

sabella Present Indistinct dorsally, triangu- Setiger 1: 3—6 long, 2—3 short lim- 
lar lobe ventrally. bates. Setiger 2-8: 5—6 long lim- 


bates. Setiger 3-7: 2 spatulates. 


in thoracic notopodia of Fabriciola acuseta. It is likely that small setal forms 
have been overlooked in thoracic and abdominal setigers of Fabriciola. 

The distinct difference between the staining pattern of the holotype in com- 
parison to paratypes might suggest sexual dimorphism. An alternative explanation 
might be changes in gland cell distribution in relation to size or age of individuals. 
The latter suggestion is unlikely since both large and small individuals stained 
similarly. Only Banse (1970) has examined staining variability using Euchone 
incolor Hartman, 1965, from different localities. He only noted staining differ- 
ences which were probably attributable to geographic variation, making no men- 
tion of noticeable differences within a single locality. 

Etymology.—The specific epithet refers to the three lobes at the anterior mid- 
dorsal collar margin. 

Distribution.—Fabriciola trilobata is known only from the type-locality. 


Fabricia infratorquata, new species 
Figs. 3d—j, 4; Table 2 


Material examined.—West Bay, Twin Cays, Belize; 30 cm depth; mat of Cau- 
lerpa verticillata on rootmat of Rhizophora mangle, some organic debris and 
fragments of Halimeda; 7 Apr 1982 (Array F202), 9 Apr 1982 (Array F203), 11 
Apr 1982 (Array F204); coll. K. Fauchald. Holotype: F204 D-4 (USNM 74644). 
Paratypes: F204 A-1, 1 specimen (USNM 74645); F204 A-2, 7 specimens (USNM 
74646); F204 A-3, 3 specimens (USNM 74647); F204 A-4, 2 specimens (USNM 
74648); F204 A-5, 1 specimen (USNM 74649); F204 B-1, 4 specimens (USNM 
74650); F204 B-2, 4 specimens (USNM 74651); F204 B-3, 2 specimens (USNM 
74652); F204 B-4, 1 specimen (USNM 74653); F204 B-5, 1 specimen (USNM 
74654); F204 C-2, 3 specimens (USNM 74655); F204 C-4, 1 specimen (USNM 


VOLUME 96, NUMBER 2 


Table 2—(Continued). 
Thoracic neurosetae 


Setiger 2—5: 6—8 uncini in dou- 


ble rows. Setiger 6-8: 5—6 un- 


cini in single rows. Main fang 
+ 1 large and | small tooth + 
3 rows of smaller teeth. 


6-12 uncini. Main fang + | 
large tooth + arc of smaller 
teeth. 


8—9 uncini in partial double 
rows. Main fang + 3 rows of 
smaller teeth. 


12-14 uncini in double rows. 
Main fang + 1| large tooth + 
1 small tooth + arc of 9 
smaller teeth. 


9-12 uncini. Main fang + 4 
rows of smaller teeth. 


Abdominal notosetae 


11-14 long-handled uncini; 7 
rows of teeth; 1-5 teeth per 
row. 


14 long-handled uncini; 5—6 
rows of teeth; 1-2 teeth per 
row. 


Long-handled uncini; 6—7 rows 
of teeth; 2—3 teeth per row. 


16-21 short-handled uncini; 9 
rows of teeth; 1-3 teeth per 
row. 


30 long-handled uncini; 9 rows 
of teeth. 


285 


Abdominal neurosetae 


1-2 long, narrow limbates. 


2—4 narrow limbates. 


3-4 narrow limbates. 


2-3 long and 1-2 short nar- 
row limbates. 


74656); F204 D-3, 4 specimens (USNM 74657); F204 D-4, 9 specimens (USNM 
74658); F204 D-5, 1 specimen (USNM 74659). Additional material: F202 A-1, 2 
specimens (USNM 74660); F202 B-1, 2 specimens (USNM 74661); F202 B-4, 1 
specimen (USNM 74662); F202 C-2, 3 specimens (USNM 74663); F202 C-4, 1 
specimen (USNM 74664); F202 D-2, 1 specimen (USNM 74665); F202 D-3, 5 
specimens (USNM 74666); F203 A-1, 1 specimen (USNM 74667); F203 A-5, 3 
specimens (USNM 74668); F203 B-3, 1 specimen (USNM 74669); F203 B-S, 1 
specimen (USNM 74670); F203 C-1, 1 specimen (USNM 74671); F203 C-2, 2 
specimens (USNM 74672); F203 C-4, 2 specimens (USNM 74673); F203 C-5, 1 
specimen (USNM 74674); F203 D-1, 1 specimen (USNM 74675); F203 D-2, 1 
specimen (USNM 74676); F203 D-4, 2 specimens (USNM 74677); F203 D-S, 1 
specimen (USNM 74678). 

Diagnosis.—Small species of Fabricia without palps. Peristomium concealed 
by collar segment. Ventral collar a distinct, anteriorly rounded triangular lobe. 
Dorsal and lateral parts of collar low, indistinct. Dorsal, longitudinal midline of 
collar segment occupied by broad groove. Collar segment divided into anterior 
and posterior parts by distinct annulation. Thoracic neuropodial uncini of setigers 
2-5 in irregular double rows, continuing as single rows in setigers 6-8; with 2 
unequal teeth above main fang, followed by series of smaller teeth. Abdominal 
uncini long-handled, with 7 rows of teeth. 

Description.—The holotype is a complete specimen with 8 thoracic and 3 ab- 
dominal setigers. Length 1.60 mm (0.56 mm comprising the branchial crown) and 
width 0.21 mm. 

A pair of semicircular branchial lobes are attached anteriorly, each with a large 
branchial heart situated dorsally (Fig. 4a). Three radioles are attached to each 
branchial lobe; they are unbranched and rounded externally. Four to 5 pairs of 


286 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 4. Fabricia infratorquata (paratype, USNM 74658): a—c, Dorsal ventral and lateral views of 
anterior end; d, Posterior end, dorsal view. 


VOLUME 96, NUMBER 2 287 


ciliated pinnules extend from the inner side of each radiole. The proximal pair 
are longest, with successive pairs becoming shorter so that all terminate near the 
distal end of the radiole. Distal ends of the radioles are blunt. Palps are absent. 

The collar segment completely conceals the peristomium. Ventrally, the collar 
is distinct, forming a thick, large triangular lobe which is rounded anteriorly (Fig. 
4b-c). The ventral surface of the ventral collar is heavily ciliated. Laterally and 
dorsally the collar is low and indistinct. The dorsal collar is divided by a longi- 
tudinal middorsal groove which extends from the anterior margin of the collar 
segment as a slight protrusion to near the segment’s posterior margin, and oc- 
cupies the middorsal ’% of the collar segment width. Adjacent to the posterior 
margin of the groove are a pair of indistinct narrow ridges, situated perpendicular 
to the groove and directed middorsally, with a narrow gap between the 2 ridges. 
The collar segment is separated into anterior and posterior halves by an annu- 
lation which is distinct ventrally and laterally, disappearing dorsally as it nears 
the middorsal groove. The anterior part of the collar segment is slightly longer 
than the posterior part, but the latter is wider. Ventrally, the posterior part is 
slightly inflated, overlapping the anterior portion of setiger |. The ventral half of 
the ventral collar and the epidermis of the posterior part of the collar segment 
contain a large number of glandular cells. 

All thoracic setigers are cylindrical, constricted at intersegmental grooves. 
Transition from the collar segment to setiger | is denoted by an abrupt increase 
in segment width. Setiger | is the shortest thoracic segment, about % the length 
of the collar segment. All subsequent thoracic setigers become longer. Setigers 
1-4 are widest, about 4 wider than the collar segment, and are wider than long. 
Setigers 5—8 only decrease slightly in width, but are longer than wide. The epi- 
dermis of setiger 1 is composed of glandular cells of about the same thickness as 
in the posterior portion of the collar segment. 

Abdominal setigers become successively shorter, with setiger 9 about /% the 
length of setiger 8 (Fig. 4d). All are wider than long, becoming slightly narrower 
posteriorly. Setigers 9-10 are cylindrical and setiger 11 is slightly dorsoventrally 
flattened. Separation between thorax and abdomen is distinct. Intersegmental 
grooves are indistinct on abdomen. Pad-like, swollen glandular areas occur lat- 
erally behind notopodial tori of setigers 9-10, and at the anterior lateral margins 
of setigers 10-11. 

Noto- and neuropodial thoracic tori are only slight swellings. Notopodia are 
located dorsolaterally in all segments. Neuropodia, absent from setiger |, are 
situated ventral to and slightly posterior to notopodia of remaining setigers. No- 
tosetae in setigers 1-8 include 3-4 superior long, nearly straight, limbate setae 
(Fig. 3d). Setigers 2 and 6-8 also have | inferior shorter limbate seta with a curved 
shaft (Fig. 3e). Setigers 3—S also possess 1-2 inferior short spatulate setae (Fig. 31). 
Notosetal fascicles are inserted obliquely. Thoracic neurosetae are gently curved 
acicular uncini (Fig. 3g). Uncini of setigers 2—5 occur in vertical, irregular double 
rows; the anterior row being slightly dorsal to the posterior row. Each row within 
a torus contains 3-4 uncini. Neurosetae in setigers 6-8 are in a single row of 5— 
6 uncini in each setiger. In lateral view, the distal end appears to have a main 
fang surmounted obliquely by a large tooth, followed by 3 smaller teeth. In frontal 
view, the main fang is surmounted by 2 pair of unequal teeth (Fig. 3h). Above 
the large tooth are 3 smaller teeth; proximal to this are two concentric arcs of 


288 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


teeth. Proximal to the distal end the shaft is slightly swollen, then tapers to a 
rounded end. 

Abdominal notopodia are distinct lateral tori on the posterior 14 of the seg- 
ments, and of the same length as thoracic neuropodia. Notosetae are long-handled 
uncini, numbering 14 in setigers 9-10 and 11 uncini in setiger 11 (Fig. 3j). All 
uncini have 7 rows of teeth; the number of teeth per row as follows (proximal to 
distal): 1+2+3+4+5+4+3. Proximal to the teeth the shaft is slightly constricted, 
then inflated proximally and terminated in a truncate base. Neurosetae originate 
just ventral to notopodia. No neuropodial tori are visible. All neurosetae are of 
one type: I—2 subequal, long-shafted limbate setae with a narrow limbation, and 
are directed dorsally (Fig. 3f). 

The pygidium is not clearly delimited from setiger 11, thus is assumed to begin 
just posterior to the notopodial tori. Anteriorly, the pygidium is of the same width 
as the adjacent segment, narrowing slightly posteriorly to a rounded end. It is 
dorsoventrally flattened. Except for the anterior margin, the entire pygidium sur- 
face is glandular. The anus is a slightly depressed, longitudinal, midventral slit. 

A pair of large roughly crescentic eyes (dorsal view) are visible just posterior 
to the intrasegmental annulation of the collar segment, on either side of the mid- 
dorsal groove. A pair of smaller, circular eyes are located laterally in the pygidi- 
um. A pair of translucent otocysts, slightly smaller than the anterior pair of eyes, 
are located dorsally in the anterior part of the collar segment on either side of 
the middorsal groove. 

The holotype and other large individuals are opaque and cream colored in 
alcohol. Small individuals tend to be more translucent. 

Individuals occupy soft, thick tubes (about 2—2.5 times as thick as the worm), 
constructed with plant and detrital material. The inner tube diameter is only 
slightly greater than the width of the worm, such that nearly the entire length of 
the individual is in contact with the tube. 

The holotype and some paratypes were stained with methyl green by the meth- 
od described above. In the holotype, the posterior portion of the collar segment 
and setiger | were uniformly stained dark green except for the fecal groove and 
tori, neither staining in any segment. Setigers 2-8 were uniformly stained lightly, 
giving a green hue. Setigers 9-11 stained much darker than adjacent setigers due 
to large stain-accepting cells and glandular areas. The pygidium was darkly stained 
as in other glandular areas. Paratypes stained similar to the holotype. Some spec- 
imens did show a greater degree of staining in setigers 5-8, resembling that of 
abdominal setigers. The ventral collar of one specimen did stain lightly, but oth- 
erwise it was similar to other individuals. 

Remarks .—Fabricia infratorquata is one of a group of Fabricia which possess 
a distinct, triangular ventral collar. Table 2 compares these species. Fabricia 
infratorquata is readily distinguished from F. sabella (Ehrenberg, 1836; see Hart- 
mann-Schroeder 1971:513) in that the latter possesses palps and all thoracic uncini 
occur in single rows. Fabricia bansei Day (1961:543) differs from F.. infratorquata 
in the arrangement and number of teeth on thoracic and abdominal uncini. Fa- 
bricia infratorquata closely resembles F. brunnea Hartman (1969:693) in that both 
have at least some thoracic uncini in double rows; the species differ in that the 
dorsal collar of the latter has a pair of small semicircular lobes and abdominal 
uncini have 2-3 teeth per row (1-5 teeth per row in F. infratorquata). Fabricia 


VOLUME 96, NUMBER 2 289 


gerdi Hartmann-Schroeder (1974:199) is also very similar to the new species in 

relation to the collar and occurrence of thoracic uncini in double rows; they differ 

in the arrangement of teeth on the thoracic and abdominal uncini. 
Etymology.—tThe specific epithet refers to the large, triangular ventral collar. 
Distribution.—Fabricia infratorquata is known only from the type locality. 


Discussion 


Fabriciola trilobata and Fabricia infratorquata both possess a character which 
has not been described in these genera: the asymmetrical arrangement of teeth 
above the main fang of thoracic uncini. Examination of uncini in a frontal view 
has not been common, probably due to difficulty in manipulating setae for ob- 
servation at such an angle. As a result a possible diagnostic character has been 
overlooked. 

Another setal characteristic of both species not commonly described is the 
number of teeth per row in abdominal uncini. Most species in which this has 
been described display a rather uniform number of teeth in each row, except in 
some cases of variation at extreme proximal and distal tooth rows. In the species 
described herein this pattern did not occur. In F. trilobata the number of teeth 
per row gradually increased in a proximal-distal direction, except for uncini from 
setiger 11, in which median tooth rows alternated with 3 and 4 per row. Fabricia 
infratorquata displayed an increasing number of teeth from proximal to distal, 
with a slight decrease in the 2 distal-most rows. A similar pattern was noted by 
Friedrich (1939) for Fabriciola baltica, with the arrangement: 
3+44+5+6+6+6+6+4+4. This pattern of variation is probably common in other 
species of both genera, but has yet to be examined. 

The majority of Fabriciola and Fabricia descriptions pay little attention to 
abdominal neurosetae; most reports refer to them as fine capillaries or ignore 
their presence. Closer attention should be given to these setae since size and 
structural differences, as seen in F. trilobata, could be of taxonomic use. 

The presence of an annulation on the collar segment of F. trilobata and F. 
infratorquata suggests the presence of an additional achaetous segment. Figures 
of Fabriciola limnicola given by Hartman (1951) depict an annulation very similar 
to that on F. trilobata. Annulations have also been illustrated by Day (1955) for 
F. capensis, by Day (1957) for F. mossambica, and by Friedrich (1939) for F. 
baltica. Figures of Fabricia show a similar annulation but it is situated further 
anteriorly, suggesting that it is simply a demarcation of the collar from the collar 
segment. In F. infratorquata this does not occur; the annulation is distinctly 
posterior to the collar, suggesting that the collar ‘“‘segment’’ is composed of two 
segments. At this time it can only be suggested that this feature be noted in future 
descriptions until such time as sectioning determines if it is an actual segment 
boundary. 


Acknowledgments 


I gratefully acknowledge Dr. Kristian Fauchald for his constant assistance, for 
providing space at the National Museum of Natural History, and making available 
the benthic samples from which the present species were described. His critical 
review of the manuscript is greatly appreciated. 


290 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Funding for this research was provided in part by Smithsonian Institution con- 
tract #SF2062530000. 


Literature Cited 


Banse, K. 1956. Beitrage zur Kenntnis der Gattungen Fabricia, Manayunkia und Fabriciola (Sa- 
bellidae, Polychaeta).—Zoologische Jahrbucher, Abteilung fiir Systematik, Okologie und Geo- 
graphie der Tiere 84:415—438. 

—.. 1957. Die Gattungen Oriopsis, Desdemona und Augeneriella (Sabellidae, Polychaeta).— 
Videnskabelige Meddelelser fra Dansk Naturhistorisk Forening, Kobenhaven 119:67—105. 

——. 1959. Fabricia acuseta n. sp., Fabriciola ghardagqa n. sp. und Oriopsis armandi (Claparede) 
aus dem Roten Meer (Sabellidae, Polychaeta).—Kieler Meeresforschungen 15:113—116. 

——.. 1970. The small species of Euchone Malmgren (Sabellidae, Polychaeta).—Proceedings of 
the Biological Society of Washington 83:387-408. 

———. 1972. Redescription of some species of Chone Kroyer and Euchone Malmgren, and three 
new species (Sabellidae, Polychaeta).—Fishery Bulletin 70:459-495. 

—. 1979. Sabellidae (Polychaeta) principally from the northwest Pacific Ocean.—Journal of the 
Fisheries Research Board of Canada 36:869-882. 

Day, J. H. 1955. The Polychaeta of South Africa. Part 3: Sedentary species from Cape shores and 
estuaries.—Journal of the Linnean Society of London 42:407-4S2. 

—. 1957. The polychaete fauna of South Africa. Part 4: New species from Natal and Mocam- 
bique.—Annals of the Natal Museum 14:59-129. 

——. 1961. The polychaete fauna of South Africa. Part 6: Sedentary species dredged off Cape 
coasts with a few new records from the shore.—Journal of the Linnean Society of London 44: 
463-560. 

——. 1967. A monograph on the Polychaeta of southern Africa. Part 2: Sedentaria.—British 
Museum of Natural History Publications 656:459-878. 

Fauchald, K. 1977. The polychaete worms. Definitions and keys to the orders, families and gen- 
era.—Natural History Museum of Los Angeles County, Science Series 28:1—190. 

Friedrich, H. 1939. Polychaeten—Studien. V—X. Zur Kenntnis einiger wenig bekannter odor neuer 
Polychaeten aus der westlichen Ostsee.—Kieler Meeresforschungen 3:362-373. 

Hartman, O. 1951. Fabricinae (feather-duster polychaetous annelids) in the Pacific.—Pacific Science 
5:379-391. 

——. 1969. Atlas of sedentariate polychaetous annelids from California.—Allan Hancock Foun- 
dation, University of Southern California, Los Angeles, 812 pp. 

Hartmann-Schroeder, G. 1971. Annelida, Borstenwurmer, Polychaeta.—Die Tierwelt Deutschlands 
58: 1-594. 

—. 1974. Teil II. Die Polychaeten des Untersuchungsgebietes. Jn G. Hartmann und G. Hart- 
mann-Schroeder, Zur Kenntnis des Eulitorals der afrikanischen Westkuste Zwischen Angola 
und Kap der Guten Hoffnung und der afrikanischen Ostkuste von Stidafrika und Mocambique 
unter besonderer Berucksichtigung der Polychaeten und Ostracoden.—Mitteilungen des Ham- 
burger Zoologisches Museum und Instittit, Erganzungsband 69:95-228. 

Monro, C. C. A. 1937. Notes ona collection of Polychaeta from South Africa, with the description 
of a new species belonging to the family Sabellidae—Annals and Magazine of Natural History 
(Series 10) 19:366-370. 


Division of Worms, Department of Invertebrate Zoology, National Museum of 
Natural History, Smithsonian Institution, Washington, D.C. 20560. Current ad- 
dress: Texas A&M University, Building 311, Fort Crockett, Galveston, Texas 
77550. 


PROC. BIOL. SOC. WASH. 
96(2), 1983, pp. 291-300 


BERMUDALANA ARUBOIDES, A NEW GENUS AND 
SPECIES OF TROGLOBITIC ISOPODA 
(CIROLANIDAE) FROM MARINE 
CAVES ON BERMUDA 


Thomas E. Bowman and Thomas M. Iliffe 


Abstract.—Bermudalana aruboides, a blind unpigmented cirolanid isopod rep- 
resenting a new genus, is described from inland marine caves on Bermuda. It is 
the first hypogean cirolanid known from Bermuda. The close resemblance be- 
tween Bermudalana and Arubolana, known from brackish groundwater on Aru- 
ba, Netherlands Antilles, suggests that both were derived from a common marine 
ancestor. 


The inland marine caves of Bermuda contain a rich and diverse invertebrate 
fauna (Sket and Iliffe 1980). The extensive nature of the underwater portions of 
these limestone caves, some being more than 1.5 km in length (Iliffe 1980), and 
the presence of strong tidal currents passing through them have resulted in the 
establishment of a considerable range of biotopes. In coastal cave entrances and 
those cave passages with strong tidal currents, sponges, bryozoans, hydroids, 
and other encrusting organisms literally cover all rock surfaces. Deeper into the 
caves, where tidal currents become more diffuse, sponges and other associated 
fauna become far less numerous. In the deepest inland caves most remote from 
the sea, the walls are totally barren of encrusting organisms and only specially 
adapted troglobitic species are present. This zonation approximately corresponds 
to that recognized in terrestrial caves (Poulson and White 1969): a twilight zone 
near the entrance, a middle zone of complete darkness and variable temperature, 
and a zone of complete darkness and constant temperature in the deep interior. 
In marine caves as well as in terrestrial caves, the twilight (coastal entrance) zone 
has the largest and most diverse fauna; the middle zone has some species which 
may commute to the surface (bats and crickets in terrestrial caves; lobsters in 
marine caves). The deep interior cave, while being the most depauperate, pos- 
sesses environmental and faunal aspects unique to caves. 

The existence of marine troglobites is a relatively new discovery. As recently 
as 1965, Vandel stated that ‘‘animals (from marine caves) have not usually under- 
gone noticeable modification’’ and thus ‘“‘marine caves . . . have but a slight in- 
terest to the biospeleologist’’ (1965:8). In our faunal survey of Bermuda’s marine 
caves, the following troglobitic species have so far been described: Atlantasellus 
cavernicolus, an isopod representing a new family (Sket 1979); Somersiella ster- 
reri and Typhlatya iliffei, two new species of caridean shrimp (Hart and Manning 
1981); and Mesonerilla propsera, a new archiannelid polychaete (Sterrer and Iliffe 
1982). Additional new species described from Bermuda’s caves which may or 
may not be troglobitic include Miostephos leamingtonensis, a new calanoid co- 
pepod (Yeatman 1980); and Apseudes bermudeus, a new hermaphroditic tanaida- 
cean (Bacescu 1980). We here describe a new genus and species of troglobitic 
cirolanid isopod from deep interior marine caves on Bermuda. 


292 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 1. Church Cave, Bermuda, seen from entrance. 


Bermudalana, new genus 


Diagnosis.—Eyes absent. Without pigment except brown incisors of mandi- 
bles. Pleonites all free, all reaching lateral margin of pleon. Frontal lamina with 
keel. Clypeus produced ventrally into slender conical process. Peduncles of an- 
tenna | and 2 3- and 5-segmented; flagella with relatively few segments. Maxilla 
2 reduced, palp and exopod lacking, endopod with only a few distal setae. Max- 
illiped with sparse setation, palp 4-segmented, endite with | retinaculum. Per- 
eopods I-3 prehensile; pereopods 4—7 slender, ambulatory. Pleopods 1-2 with 
undivided setose rami. Pleopods 3—5 with 2-segmented setose exopods and un- 
divided endopods; endopods of pleopods 3-4 with a few apical setae, endopod 
of pleopod 5 without setae. Appendix masculina inserted subterminally. Penes 
well developed, narrowly cylindrical, separated at base. 

Type species.—Bermudalana aruboides, new species. 

Etymology.—From the locality, Bermuda, plus (Ciro)lana. Gender feminine. 


Bermudalana aruboides, new species 
Figs. 2-4 


Material.—Bermuda: Church Cave (also known as Paynter’s Vale Cave), Ham- 
ilton Parish, 11) July 1982, leg. 1. M. Iiife, 4 6 G95 3°85 3-6; 321 inm)yandene 
(4.1, 3.3 mm) were collected from 7 to 10 m water depths with scuba using a 
suction bottle. Wonderland Cave (also known as Whitby Cave), Hamilton Parish, 
19 May 1982, leg. T. M. Iliffe, 2 ¢ (3.7, 3.4 mm) and 3 @ (4.0, 4.0, ? mm) were 
collected from 10 to 15 m depths with scuba using a suction bottle. The 3.9 mm 
36 from Church Cave is the holotype (USNM 195020); the other specimens are 


VOLUME 96, NUMBER 2 293 


paratypes (Church Cave specimens USNM 195021, Wonderland Cave specimens 
USNM 195022). 

Distribution.—Known only from the anchialine habitats of Church and Won- 
derland Caves, Bermuda. 

Habitat.—The Bermuda islands consist of a mid-ocean volcanic seamount 
capped with Pleistocene and Recent, marine and eolian limestones. Bermuda is 
one of the world’s most geographically isolated islands, located 1000 km east of 
the North American continent in the section of the North Atlantic known as the 
Sargasso Sea. Bermuda has never been part of a continental land mass. The 
island’s limestone caves were formed during low stands of sea level corresponding 
to periods of Pleistocene glaciation (Bretz 1960; Palmer et al. 1977; Iliffe 1981). 
As postglacial sea levels rose, much of the former extent of the caves was flooded 
by sea water. Approximately 200 inland caves are known from Bermuda, over 
half of which contain tidal, sea level pools. 

Church Cave is located on the grounds of the Castle Harbour Hotel about 250 
m linear distance from Castle Harbour, the nearest body of open salt water. The 
main entrance is about 26 m above sea level and consists of an opening 25 m 
wide by 15 m high in a collapse sink 25 m in diameter. Inside this entrance, a 
long steep breakdown slope descends to a sea level lake 35 m wide by 35 m across 
(Fig. 1). A small amount of sunlight from the entrance can reach one corner of 
the lake, but the rest remains in total darkness. The tides in this lake have an 
average range of 40% that of the open sea and have a lag time of about 107 
minutes. Surface salinity is between 15.5 and 22.8%c, probably varying with rain- 
fall, while at 1, 10, and 20 m depths, salinities are about 27, 34.5, and 35.3%o 
respectively. Surface temperatures seasonally range from about 16 to 19°C, but 
temperatures in deeper waters (20.3°C at 20 m) remain nearly constant year round. 
The natural geothermal gradient has been proposed as a possible explanation for 
this anomalous temperature increase with depth (lliffe et a/. 1983). The sides 
of the lake are undercut, with the bottom being 20 m at the deepest point and 
floored with breakdown blocks of considerable size, but little fine sediments. The 
underwater parts of the cave are abundantly decorated with delicate speleothems, 
including “‘soda straws’’ and helectites, all perfectly preserved despite their long 
submersion. The presence of such speleothems, which only form in air, in all 
explored parts of the underwater caves indicates that the caves were dry during 
the prolonged periods of Pleistocene glaciation when sea levels were 80 to 100 m 
lower than today. Only one underwater cave passage has been found extending 
away from the lake and this ends after 45 m in a flowstone plug. 

Wonderland Cave is located 2 km northwest of Church Cave and is 420 m from 
Castle Harbour. It was once operated as a commercial cave, but has not been 
used for such purposes since the 1940s. A steep set of 89 steps descends from 
the small entrance building located at an elevation of 24 m to a large room de- 
veloped along a linear inclined fissure. The room contains a sea level lake 60 m 
long by as much as 12 m wide. This lake is 18 m at the deepest point and floored 
by large slabs of breakdown. A 50 m long underwater passage extends from the 
far side of the lake to re-emerge in an air-filled breakdown room. No other un- 
derwater passages have been found. As in Church Cave, many large stalactites 
and stalagmites are present even in the deepest parts of the lake. 

All specimens of Bermudalana were taken from open waters of the lakes by 


294 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


VOLUME 96, NUMBER 2 29 


Nn 


cave divers as the animals were observed swimming several meters above the 
bottom. Divers used a bright underwater light to scan the very clear cave waters. 
Any animals or particles in the water would flash as the light beam passed across 
them, effectively locating them. The swimming behavior of Bermudalana is most 
likely a result of the animal’s food locating actions and not an escape reaction. 
No specimens were ever observed crawling over the substrate. The lack of any 
encrusting organisms with only planktonic or nektonic species being observed 
also supports the idea that Bermudalana is a predator, capturing its food from 
open waters. Other animals collected or observed from Church and Wonderland 
Caves include a halocyprid ostracod now under study, a peracarid representing 
a new order (Bowman and Iliffe, in preparation), and a caridean shrimp, probably 
Typhlatya iliffei. Copepods are probably also present as they have been found in 
nearly every marine cave studied in Bermuda. It is likely that Bermudalana will 
also be found in other far inland caves of Bermuda as they are investigated. 

Description.—Body moderately slender, slightly more than 3 x as long as wide, 
length about 4 mm. Anterior margin of head slightly concave on either side of 
minute rostrum, about 1.7x as wide as long. Frontal lamina visible in dorsal 
view, about a third longer than wide, ventral surface produced into carina. Clyp- 
eus in lateral view produced into rather slender cylindrical process. 

Pereonites | and 5—7 subequal in length, distinctly longer than the subequal 
pereonites 2—4; all pereonites with rounded posteroventral corners. Posteroven- 
tral corners of coxae 2—4 rounded, of 5—7 with small points. Pleon (excluding 
telson) about half length of pereon; pleonites 1-4 subequal in length, pleonite 5 
shorter and slightly narrower, epimera all ending in small points. Telson lingui- 
form, slightly shorter than width at base, posterior sixth with 4 setae on each 
side set in marginal notches; marginal spines absent. 

Antenna | reaching slightly beyond posterior margin of pereonite 1; peduncle 
segment 3 very long, peduncle segments 2-3 with long plumose distal setae; 
flagellum 5-segmented, Ist segment nearly 3x as long as remaining segments 
combined, all segments with long esthetes. Antenna 2 reaching posterior margin 
of pereonite 6; segments of peduncle successively longer; flagellum 8-segmented. 

Incisors of mandibles with 3 cusps, cusps more deeply separated in right man- 
dible; left lacinia with 10 spines, right lacinia with 9 spines; molar with 14 spines; 
segment 2 of palp about 2.5x< as long as segment 1, with about 7 marginal setae 
on distal fifth; segment 3 slightly shorter than segment |, with about 7 setae. 

Maxilla | exopod apex with 10 spines, 2 much longer than others, and a central 
seta; endopod with 3 apical spines and 2 subterminal setae. Maxilla 2 reduced to 
single ramus armed apically with 4 long and 2 short setae. Maxilliped endite with 
2 apical plumose setae. 

Pereopods !—3 subchelate, dactyl closing against palm of expanded propus; 
palm bearing distally a spine with posterior setule, 3-5 thickset spines, and 8-15 


— 


Fig. 2. Bermudalana aruboides: A, Habitus, dorsal; B, Coxae and epimera, lateral; C, Pleon and 
telson, dorsal; D, Antenna 1; E, Antenna 2 peduncle; F, Frontal process, clypeus, and labrum; G, 
Clypeus, lateral; H, Left mandible; I, Incisor, lacinia, and molar of right mandible; J, Maxilla 1; kK, 
Maxilla 1 exopod, apex; L, Maxilla 2. 


296 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 3. Bermudalana aruboides: A, Maxilliped; B, Maxilliped endite; C-I, Pereopods 1-7. 


VOLUME 96, NUMBER 2 297 


Fig. 4. Bermudalana aruboides: A—B, Pleopods 1-2, 2; C, Pleopod 2 endopod, 5; D-F, Pleopods 
3-5, 2; G, Penes, 3. 


more slender spines; posterior margin of carpus with 4—7 slender spines. Propus 
of pereopod 2 slightly less expanded but longer than that of pereopod 1; propus 
of pereopod 3 slender. Pereopods 4—7 slender, sparsely armed with spines, mostly 
at distal ends of segments; basipods with long setae on posterior margins, 2 on 
pereopods 4—5, 1 on pereopods 6-7. 


298 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Exopods of pleopods 1-4 with long marginal setae, some of those on apex of 
pleopods 3-4 longer, thicker, and ending in terminal setule. Exopod of pleopod 
5 with 4 medial and | lateral setae on distal segment; apex without setae. En- 
dopods of pleopods 3-4 much narrower than very broad exopods, armed with 2 
apical setae in pleopod 3, 3 in pleopod 4; endopod of pleopod 5 without setae. 
Appendix masculina of d pleopod 2 about 0.6x as long as endopod, with con- 
striction near base, curving slightly laterally and ending acutely. 

Uropod endopod pyriform, about a third longer and 3x as wide as sublinear 
exopod; medial margin and apex armed with long setae, lateral margin with 2 
short setae in distal part and 2 long setae set in from margin slightly distal to 
midlength. Exopod with 2 long medial setae, 2 short lateral setae, and cluster of 
setae lateral to apex, medial part of which is produced into narrow process. 

Etymology.—The specific name refers to the important similarities between 
Bermudalana and Arubolana. 

Relationships.—TYhe combination of five pleonites reaching the lateral margin 
of the pleon and pereopods 1-3 prehensile is found in two genera of Cirolanidae, 
Bahalana Carpenter, 1981, with one species from Lighthouse Cave, San Salva- 
dore Island, Bahamas, and Sphaeromides Dollfus, 1897, with three species from 
caves adjacent to the Mediterranean. These genera differ from Bermudalana in 
having a normally developed maxilla 2, a 5-segmented palp on the maxilliped, 
and a basally inserted appendix masculina. 

The closest relative of Bermudalana is Arubolana Botosaneanu and Stock, 
1979, represented by a single species, A. inula Botosaneanu and Stock, from 
brackish groundwater in Aruba, Netherlands Antilles. The two genera have in 
common reduced segmentation of antennae | and 2, a greatly reduced maxilla 2, 
a maxilliped with a 4-segmented palp, an appendix masculina inserted subter- 
minally, and interrupted marginal setae on the exopod of pleopod 5. Arubolana 
differs in having pleonite 5 overlapped laterally by pleonite 4 and only pereopods 
1-2 prehensile. These differences are usually believed to be of generic value in 
the Cirolanidae, hence we have proposed a new genus for the Bermuda cirolanid. 
We realize, however, that a plausible case could be made for assigning the latter 
to Arubolana or for recognizing Bermudalana as a subgenus of Arubolana. 

Origin.—The common possession by Arubolana and Bermudalana of several 
specialized character-states not found elsewhere in the Cirolanidae can be ex- 
plained best by their evolution from a common ancestor. Convergent evolution 
could be evoked for a single character-state but is highly unlikely for such a 
combination of several character-states. Aruba and Bermuda are separated by 
about 1400 miles (2250 km) and water of abyssal depths; dispersal from one of 
these islands to the other by a small troglobitic isopod would seem to be a remote 
possibility. Rafting on floating objects carried from Aruba to Bermuda via the 
Gulf Stream cannot be disproven, but we do not consider it a serious possibility. 
The origin of the two species from a common marine ancestral species is a more 
believable alternative, but two difficulties must be faced. 1. Cirolanids are rarely 
found at the abyssal depths (>2000 m) of most of the Atlantic between Aruba 
and Bermuda. Depths recorded for three blind deep-sea species of Cirolana are 
C. caeca Dollfus, 1903, 1200-2500 m; C. anocula Kensley, in press, 750 m; C. 
californiensis Schultz, 1966, 700-2000 m (Brusca and Ninos 1978). 2. Prehensile 
pereopods in the Cirolanidae are known only in hypogean species. The reason 


VOLUME 96, NUMBER 2 299 


for this is not known, but might be related to a shift in feeding from scavenging 
to predation in cave cirolanids. Carpenter (1981) reports that Bahalana geracei 
holds its prey firmly with prehensile pereopods !-3 and bites off small pieces 
with its mandibles. 

Nevertheless, Aruba and Bermuda have never been connected by land and if 
Arubolana and Bermudalana had a common ancestor, as we firmly believe, it 
must have been a marine cirolanid. It is possible that the ancestral cirolanid lacked 
prehensile pereopods and that the latter evolved independently in the two genera 
after reaching their present localities. 


Acknowledgments 


This study was supported by a National Science Foundation Grant (DEB- 
8001836) to Thomas M. Iliffe. We thank Drs. C. W. Hart, Jr., and R. B. Manning 
for their reviews of the manuscript. Cave diving equipment and techniques used 
to conduct this study met standards of the National Speleological Society. This 
paper is Contribution No. 939 from the Bermuda Biological Station for Research, 
ie: 


Literature Cited 


Bacescu, M. 1980. Apseudes bermudeus n. sp. from caves around the Bermuda Islands.—Acta 
Adriatica 21:401—407. 

Botosaneanu, L., and J. H. Stock. 1979. Arubolana imula, n. gen., n. sp., the first hypogean cirolanid 
isopod found in the Lesser Antilles.—Bijdragen tot de Dierkunde 49(2):227-233. 

Bretz, J. H. 1960. Bermuda: A partially drowned, late mature, Pleistocene karst.—Bulletin of the 
Geological Society of America 71:1729-1754. 

Brusca, R. C., and M. Ninos. 1978. The status of Cirolana californiensis Shultz, and C. deminuta 
Menzies and George, with a key to the California species of Cirolana (Isopoda: Cirolanidae).— 
Proceedings of the Biological Society of Washington 91(2):379-385. 

Carpenter, J. H. 1981. Bahalana geracei n. gen., n. sp., a troglobitic marine cirolanid isopod from 
Lighthouse Cave, San Salvador Island, Bahamas.—Bijdragen tot de Dierkunde 51(2):259-267. 

Dollfus, A. 1897. Sur deux types nouveaux de Crustacés Isopodes appartenant a la faune souterraine 
des Cevennes.—Comptes Rendus Hebdomadaires des Séances de |’Académie des Sciences, 
Paris 125:130-131. 

———. 1903. Note préliminaire sur les especes de genre Cirolana recuillies pendent les campagnes 
de ‘l’Hirondelle’ et de la “Princesse Alice’.—Bulletin de la Société Zoologique de France 28(2): 
5-10. 

Hart, C. W., Jr., and R. B. Manning. 1981. The cavernicolous caridean shrimps of Bermuda (AI- 
pheidae, Hippolytidae, and Atyidae).—Journal of Crustacean Biology 1(3):441-456. 

Iliffe, T. M. 1980. Mid-ocean cave diving.—Underwater Speleology 7(4):46—-48. 

1981. The submarine caves of Bermuda.—Proceedings of the Eighth International Congress 

of Speleology, Bowling Green, Kentucky, U.S.A., pp. 161-163. 

——., C. W. Hart, Jr., and R. B. Manning. 1983. Biogeography and the caves of Bermuda.— 
Nature 302:141-142. 

Kensley, B. In press. The South African Museum’s Meiring Naude cruises. Marine Isopoda of 
the 1977, 1978 and 1979 Cruises.—Annals of the South African Museum. 

Palmer, A. N., M. V. Palmer, and J. M. Queen. 1977. Geology and origin of the caves of Bermuda.— 
Proceedings of the Seventh International Congress of Speleology, Sheffield, U.K., pp. 336— 
339. 

Poulson, T. L., and W. B. White. 1969. The cave environment.—Science 165:971—981. 

Sket, B. 1979. Atlantasellus cavernicolus n. gen., n. sp. (Isopoda Asellota, Atlantasellidae n. fam.) 
from Bermuda.—Bioloski Vestnik (Ljubljana) 27(2):175-183. 


300 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


, and T. M. Iliffe. 1980. Cave fauna of Bermuda.—Internationale Revue der gesamten Hy- 

drobiologie 65(6):87 1-882. 

Sterrer, W., and T. M. Iliffe. 1982. Mesonerilla prospera, a new archiannelid from marine caves 
on Bermuda.—Proceedings of the Biological Society of Washington 95(3):509-514. 

Vandel, A. 1965. Biospeleology: The biology of cavernicolous animals.—Pergamon Press, Oxford, 
U.K., 524 pp. 

Yeatman, H. C. 1980. Miostephos leamingtonensis, a new species of copepod from Bermuda. — 

Journal of the Tennessee Academy of Science 55(1):20-21. 


(TEB) Department of Invertebrate Zoology (Crustacea), NHB 163, Smithson- 
ian Institution, Washington, D.C. 20560; (TMI) Bermuda Biological Station for 
Research, Ferry Reach I-15, Bermuda. 


PROC. BIOL. SOC. WASH. 
96(2), 1983, pp. 301-306 


VARICHAETADRILUS, A NEW NAME FOR VARICHAETA 
BRINKHURST, 1981, NON SPEISER, 1903, 
(DIPTERA) WITH A DESCRIPTION OF A 

NEW SPECIES V. FULLERI 


Ralph O. Brinkhurst and R. Deedee Kathman 


Abstract.—The genus Varichaeta Brinkhurst, 1981, is renamed Varichaetad- 
rilus. Varichaetadrilus fulleri, new species, is described from material collected 
in Kentucky. The species has distinctive penes and lacks dorsal hair and pectinate 
setae; all setae are bifid. The bifidus form of V. pacificus is described. It differs 
from the typical form by the lack of hair and pectinate setae. 


The genus Varichaeta Brinkhurst, 1981, was erected because a clear distinction 
could be made between the male ducts of this genus and those of Isochaetides 
Hrabé, 1966 (Brinkhurst 1981). Two of the previous Varichaeta species had been 
assigned to Jsochaetides (or its precursor I[sochaeta Pointner, 1911; see Brink- 
hurst 1981 for clarification) as all of the species involved have very long, tubular 
male ducts. In Isochaetides the vasa deferentia are very long and the atria are 
quite short, but in Varichaeta the atria are at least as long as the vasa deferentia. 
The penes in Varichaeta are unusually large, erectile, and have short cuticular 
sheaths at the distal end only. The Isochaeta—Isochaetides complex belongs to 
those few genera in which all the species lack dorsal hair and pectinate setae 
(i.e., Clitellio Savigny, 1820, and Limnodrilus Claparede, 1862) which is the basis 
for the name. Other genera (Potamothrix Vejdovsky and Mrazek, Aulodrilus 
Bretscher for example), contain species with hair and pectinate setae and others 
that lack them, but there has never been any suggestion of splitting these genera 
on the basis of setal pattern. In fact, this degree of variation is given only sub- 
specific or varietal status by several authors in the absence of any other significant 
differences in the taxa concerned (see Tubifex tubifex (Muller, 1774); Ilyodrilus 
frantzi Brinkhurst, 1965; Potamothrix prespaensis (Hrabé, 1931)—all in Brink- 
hurst 1971). 

While the male ducts of V. nevadana (Brinkhurst, 1965) and V. israelis (Brink- 
hurst, 1971) were thought to be of the form found in /sochaeta—Isochaetides, the 
inclusion of both species in that complex was therefore acceptable despite the 
presence of hair and pectinate setae. Once the male ducts of these two plus V. 
pacifica Brinkhurst, 1981, could be recognized as distinctive, the genera were 
separated by Brinkhurst (1981). The name Varichaeta was used to indicate the 
presence of hair and pectinate setae in contrast to Isochaeta. As the name Var- 
ichaeta proves to be preoccupied (Varichaeta Speiser, 1903—Diptera) we now 
propose the name Varichaetadrilus for the three known species, Varichaetadrilus 
pacificus (Brinkhurst, 1981) new combination (type-species), Varichaetadrilus 
nevadana (Brinkhurst, 1965) new combination, and Varichaetadrilus israelis 
(Brinkhurst, 1971) new combination. 

In addition, specimens collected by S. L. H. Fuller (Academy of Natural Sci- 


302 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


ences, Philadelphia) in Kentucky prove to belong to this genus, and these will 
now be described. 


Varichaetadrilus fulleri, new species 
Fig. 1 


Diagnosis.—Worms up to 50 mm long, about | mm wide anteriorly, more than 
200 segments. Prostomium small, pharynx large and eversible, septa in anterior 
segments very thick, muscular. Setae bifid, 2-3 or up to 5 per bundle anteriorly, 
1, sometimes 2 from behind the clitellum; upper teeth shorter than the lower in 
1 or 2 anterior segments, longer than the lower in most preclitellar bundles (or 
worn and broken to appear shorter); upper teeth shorter and thinner than the 
lower posteriorly. No genital setae. Spermathecal pores in position of missing 
ventral setae of X, male pores slightly median to line of (missing) ventral setae 
of XI, female pores in the same line in 11/12. Spermathecae voluminous with 
elongate spermatozeugma; ducts wide, capacious with folded walls and cervix- 
like plugs between ampullae and ducts. Male ducts exceedingly long, both vasa 
deferentia and atria elongate; prostate glands small, attached to atria close to 
union with vasa deferentia; penes very large, erectile, with cuticular sheaths on 
distal ends only; terminal parts of ejaculatory ducts make S-bends before entering 
penes. 

Material examined.—Holotype: USNM 79466, | dissected mature specimen 
on 2 slides, Canada Balsam preparation. Type-locality: 3.2 miles WNW of Birk 
City, Daviess Co., Kentucky, 11 July 1981. Paratypes: 2 mature specimens, Aca- 
demy of Natural Sciences, Philadelphia, coll. 3.2 miles WNW of Birk City, Davies 
Co., 30 July 1981; USNM 79467-79475, 8 mature and 6 immature specimens on 
12 slides; Brinkhurst collection: 1 mature specimen on | slide; Kathman collec- 
tion: | mature specimen on | slide; localities as above plus other localities near 
Birk City and Green River 3.5 miles SE of Calhoun, McLean County, Kentucky, 
all coll. S. L. H. Fuller, May—July 1981. 

Etymology.—‘‘fulleri’’—for S. L. H. Fuller, the collector. 

Remarks.—The enormously elongate male ducts are impossible to illustrate in 
their entirety as they cannot be dissected out in one piece or seen in whole mounts 
with ease, but they resemble those of the other species in the genus. The relative 
length of each vas deferens cannot be determined but the recognition of several 
fragments of the vas suggests it may be as long as the atrium. The penes are 
erectile and are seen in various degrees of retraction in the material available. 
The setae are all bifid, unlike those of the other species in the genus, but this 
does not exclude this new species as other well-established genera share this 
variation (see above for some of many examples). The large size of these worms 
may be a product of local environment, as specimens of Limnodrilus hoffmeisteri 
Clap., from the same localities are unusually large. A total of 67 samples which 
contained oligochaetes were examined in this survey, executed in three searches 
in 1981 (May-June, July-August, and October). The new species was found in 
association with other tubificids (Limnodrilus hoffmeisteri 18 samples, L. mau- 
meensis 12, L. udekemianus 9, L. cervix 2, Branchiura sowerbyi 15) and on one 
occasion, with another unidentifiable large tubificid with bifid setae. Other tubi- 
ficids were found in samples not containing Varichaetadrilus (Limnodrilus cla- 
paredeianus, L. angustipenis, Aulodrilus pigueti) and it was not associated with 


VOLUME 96, NUMBER 2 303 


Fig. 1. Varichaetadrilus fulleri (from type-series): A, Spermatheca showing valve at union of duct 
with ampulla (s = spermatozeugma); B, Four pieces of male duct (a = atrium, f = funnel, p = pros- 
tate, v = vas deferens); C, Penis in sigmoid form of the in situ condition (teased apart in B); D, setae. 


304 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


the naidids Dero digitata, Nais variabilis, N. pardalis, N. bretscheri and Hae- 
monais waldvogeli or the large worm Sparganophilus eiseni. The new species 
appears to be very tolerant of organic pollution, being found in association with 
an indicator assemblage. 

Other collections made by the authors from the Green River and its tributaries 
in the Mammoth Cave system (well upstream of the localities cited here) failed 
to reveal this new species, and so the discovery of V. fulleri in the Green River 
does not appear to be due to downward migration from a refuge cave environ- 
ment. As we know of no unique characteristic of the Green River, we can only 
assume that V. fulleri has been overlooked or mistaken for a Limnodrilus species 
until now, and that it is, in fact, widely distributed. However, the senior author 
failed to find it in his examination of all of the collections of the Academy of 
Natural Sciences, Philadelphia, from the rivers along the southeast seaboard of 
the U.S.A. up to 1962, and the junior author has similarly identified material from 
detailed studies in Tennessee, Alabama, Georgia, and some sites in South Car- 
olina without seeing it, and it has not been reported by scientists actively working 
with oligochaetes in Louisiana. It is possible that it forms part of a lower Mis- 
sissippi fauna, a suggestion confirmed by the recent independent discovery of the 
species by C. R. Bingham (personal communication), who will describe that ma- 
terial elsewhere. 


Varichaetadrilus pacificus (Brinkhurst, 1981) 
Fig. 2 


Seven specimens of this species were found in the Columbia River at Snag 
Island (lower elevation, samples A and E, August 1980, Miller Sands area) by 
Dr. R. J. Diaz from samples collected by the U.S. Corps of Engineers. All seven 
lack hair and pectinate setae, and are therefore termed the bifidus form of the 
species. One other fragment has a few sparse hair setae and may also belong to 
this species, but that could not be determined. 

This finding further substantiates the belief that the presence of hair and pec- 
tinate setae can probably be affected by environmental conditions. This site is 
subjected to occasional intrusions of salt water. The senior author has also seen 
specimens of the bergi and blanchardi forms of T. tubifex from springs 70 km 
from Tripoli, Libya (Dr. C. S. Woods, personal communication), but the bergi 
forms had hair setae in postclitellar bundles only. Such sites are well known to 
have elevated conductivity levels in the water. The blanchardi form was recently 
reported from France by Giani and Martinez-Ansemil (1981), in the Eau Salée, 
which descriptive name alone makes the point. 

Poddubnaya (1980) obtained new material of the bergi form from the type- 
locality (Lake Issyk-Kul). She stated that the presence of the typical form and 
the variant form in the same locality precludes the possibility of regarding these 
as subspecies, as discussed earlier by Brinkhurst (1971) who reduced these vari- 
ants to ‘‘forms’’ and who has subsequently used this terminology consistently 
for this same setal variation where other characteristics do not differ. Poddubnaya 
presents measurements of 14 characteristics of the male ducts and spermathecae, 
giving error terms (the number of specimens measured is 100 or more of each 


VOLUME 96, NUMBER 2 305 


Fig. 2. Varichaetadrilus pacificus, bifidus form: A, Male duct from funnel to penis, fragmented; 
B, Penis with penial setae. 


form). These values establish the fact that the size of these parts may be smaller 
(vas deferens parameters) or larger (atrial parameters) in the bergi form in com- 
parison to the tubifex form, but the form of the parts is quite similar. The only 
major distinction (in our opinion) may be the gradual transition between the 
narrow and wide parts of the longer vas deferens in tubifex as opposed to the 
gradual transition in bergi, though Poddubnaya also describes differences in the 
positions of pharyngeal glands and postseptale of the anterior nephridia, char- 
acters that are difficult to evaluate as they are so seldom described. 

The final resolution of this problem of potential setal variation within a species 
will, of course, only be solved if the bergi and blanchardi forms of T. tubifex (or 
one of the other species showing this same set of variations) can be produced by 
culturing experiments. 

The reference to setae in the generic definition should be omitted. This char- 
acteristic should only be used in those genera in which the hair setae are char- 
acteristically absent. 


Acknowledgments 


We wish to acknowledge Mr. S. L. H. Fuller and the Academy of Natural 
Sciences, Philadelphia, for permission to publish this description. 

The senior author is indebted to R. W. Huddleson (Chevron Oil Field Research 
Co.) for noting the nomenclatural problem in his assiduous monitoring of the 
literature, and to the late H. R. Baker for assistance. The manuscript was prepared 
by M. Stone. 


Literature Cited 


Brinkhurst, R. O. 1971. Jn Brinkhurst, R. O., and B. G. M. Jamieson. Aquatic oligochaeta of the 
world.—Oliver and Boyd, Edinburgh, xi + 806 pp. Part 2. Systematics. 8. Family Tubificidae, 
pp. 444-625. 

——. 1981. A contribution to the taxonomy of the Tubificinae (Oligochaeta: Tubificidae).—Pro- 
ceedings of the Biological Society of Washington 94: 1048-1067. 

Giani, N., and E. Martinez-Ansemil. 1981. Contribution a la connaissance des oligochaetes aqua- 
tiques du bassin de l’Argens (Var, France).—Annales de Limnologie 17:121-141. 


306 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Poddubnaya, T. L. 1980. Independence of the species Tubifex bergi (Hrabé) (Oligochaeta, Tubifi- 
cidae) from Lake Issyk-Kul.—Trudy Instituta Biologit Vnutrennikh vod Akademii Nauk SSSR 
41:41-S2. 


(ROB) Ocean Ecology Laboratory, Institute of Ocean Sciences, P.O. Box 6000, 
Sidney, British Columbia V8L 4B2, Canada. (RDK) E.V.S. Consultants Ltd., 
Box 8, Marine Technology Center, Sidney, British Columbia V8L 3S1, Canada. 


PROC. BIOL. SOC. WASH. 
96(2), 1983, pp. 307-308 


BIOLUMINESCENCE IN THE MARINE OSTRACOD 
CYPRIDINA AMERICANA (MULLER, 1890) 
OFF MANZANILLO, MEXICO 
(MYODOCOPA: CYPRIDININAE) 


David Lapota 


Abstract.—A myodocopid ostracod, Cypridina americana (Miller, 1890), found 
in coastal waters off Manzanillo, Mexico, was observed to be bioluminescent. 
Collection of this ostracod extends the known geographic distribution of this little- 
studied species and documents the only observation of bioluminescence in a 
species of Cypridina found in Eastern Pacific coastal waters. 


Marine ostracods create spectacular bioluminescent displays in the world’s 
oceans (Haneda 1940; Harvey 1952; Turner 1965; Tett and Kelly 1973). Specifi- 
cally, luminescence has been observed in five species of Vargula (Kornicker and 
King 1965; Seliger and McElroy 1965; Shimomura et al. 1969; Raymond and 
DeVries 1976; Kornicker and Baker 1977; Morin and Bermingham 1980), one 
species of Pyrocypris (Tett and Kelly 1973), 22 species of Conchoecia (Rudjakov 
1967; Angel 1968), and two species of Cypridina (Haneda 1940, 1953; Tsuji et al. 
1970). 

During the Varifront III cruise (16 Nov to 16 Dec 1981) into the Gulf of Cali- 
fornia, a third species of Cypridina was observed to be bioluminescent. The 
details of this observation are reported herein. 

A plankton net (35 wm mesh, | m long) was towed at the sea surface from the 
USNS DeSteiguer at approximately two knots for 15 minutes on 27 November, 
1800 hrs (GMT 331, 0100 hrs) at 19°23.8’N, 105°18.9’W, one hour prior to sunset. 
The sample was diluted with freshly filtered (0.45 wm) seawater for isolation of 
organisms and subsequent observation of bioluminescence. Only two ostracods, 
both actively swimming, were found within the entire collection. After several 
washings, each was placed into an individual vial. One of the two specimens, an 
adult male, was identified as Cypridina americana (Muller, 1890) by Dr. Louis 
S. Kornicker and Anne C. Cohen of the Smithsonian Institution and is deposited 
in the U.S. National Museum (USNM 159080). 

Three hours following isolation of the ostracods, the vials were inadvertently 
jarred and a point of blue light was immediately observed within each ostracod. 
A second jarring of the vials produced an intense and persistent blue luminous 
cloud from each. Within the clouds were points of light that persisted for at least 
one minute. Subsequent jarring of the ostracods produced only brief and some- 
times delayed flashes observable for only several seconds. The source of the 
discharged luminous clouds from these ostracods was not investigated although 
the mechanism for light production in the ostracod Vargula hilgendorfii (Muller, 
1890) has been well documented (Harvey 1916, 1952). 

Miller (1890) described Cypridina americana from specimens collected off the 
west coast of Colombia and Ecuador at S5°N 82°W, 3°N 85°W, and 3°S 81°W. 
Cypridina americana was reportedly found off Hawaii (Sharpe 1908) but Muller 


308 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


(1912) identified the specimen as Cypridina sharpei. Other than Miiller’s (1890) 
description of C. americana, no other reports were found in the literature (per- 
sonal communication, L. S. Kornicker). Collection of this ostracod off Manzani- 
llo, Mexico extends the known geographic distribution of this little studied species 
and also documents the only observation of bioluminescence in a species of 
Cypridina in Eastern Pacific coastal waters. 


Acknowledgments 


I am indebted to Dr. Louis C. Kornicker and Anne C. Cohen, National Museum 
of Natural History, Smithsonian Institution, for identification of the luminous 
ostracod, for making available to me Miller’s collection data of the known dis- 
tribution of C. americana, and drawing my attention to Morin and Bermingham’s 
observations on V. bullae. I am also grateful for comments and suggestions from 
Drs. Kornicker and Richard V. Lynch, III (Naval Research Laboratory, Wash- 
ington, D.C.). Financial support is gratefully acknowledged from the Naval Ocean 
Systems Center Independent Research Program. 


Literature Cited 


Angel, M. V. 1968. Bioluminescence in planktonic halocyprid ostracods.—Journal of the Marine 
Biological Association, United Kingdom 48:255—257. 

Haneda, Y. 1940. Phenomenon of bioluminescence.—Seingaku-Shidoshu 5:18-31. 

1953. Observation on some marine luminous organisms of Hachijo Island, Japan.—Recent 
Oceanographic Works of Japan 1:103—-108. 

Harvey, E. N. 1916. The mechanism of light production in animals.—Science 44:208—209. 

1952. Bioluminescence. Academic Press, New York, 649 pp. 

Kornicker, L. S., and C. E. King. 1965. A new species of luminescent Ostracoda from Jamaica, 

West Indies.—Micropaleontology 11(1):105—110. 

, and J. H. Baker. 1977. Vargula tsujii, a new species of Ostracoda from Lower and southern 

California (Myodocopa: Cypridininae) .—Proceedings of the Biological Society of Washington 

90(2):218-231. 

Morin, J. G., and E. L. Bermingham. 1980. Bioluminescent patterns in a tropical ostracod.— 
American Zoologist 20:851 (Abstract). 

Muller,G. W. 1890. Neue Cypridiniden.—Zoologische Jahrbticher, Abteilungen fiir Systematik 5(2): 

DIN—PS2, 

1912. Ostracoda.—Das Tierreich 31:1—434. 

Raymond, J. A., and A. L. DeVries. 1976. Bioluminescence in McMurdo Sound, Antarctica.— 
Limnology and Oceanography 21:599-602. 

Rudjakov, J. A. 1967. The study of the luminescence of pelagic ostracoda. Pp. 52-62, in: Bioen- 
ergetics and biological spectrophotometry.—Moscow: Nauka. 

Seliger, H. H., and W. D. McElroy. 1965. Light: Physical and biological action.——Academic Press, 
New York, 417 pp. 

Sharpe, R. W. 1908. A further report on the Ostracoda of the United States National Museum.— 
Proceedings of the U.S. National Museum 35 (1651):399-430. 

Shimomura, O., F. H. Johnson, and T. Masugi. 1969. Cypridina bioluminescence: Light emitting 
oxyluciferin-luciferase complex.—Science 164:1299-1300. 

Tett, P. B., and M. G. Kelly. 1973. Marine bioluminescence.—Oceanography and Marine Biology 
Annual Review 11:89-173. 

Tsuji, F. I., R. V. Lynch, III, and Y. Haneda. 1970. Studies on the bioluminescence of the marine 
ostracod crustacean Cypridina serrata.—Biological Bulletin 139:386—401. 

Turner, R. J. 1965. Notes on the nature and occurrence of marine bioluminescent phenomena.— 
N.I.O. Internal Report, No. 134, 30 pp. 


Radiation Physics Division, Code 534, Naval Ocean Systems Center, San Die- 
go, California 92152. 


PROC. BIOL. SOC. WASH. 
96(2), 1983, pp. 309-316 


ESCHMEYER NEXUS, A NEW GENUS AND SPECIES OF 
SCORPAENID FISH FROM FIJI 


Stuart G. Poss and Victor G. Springer 


Abstract.—Eschmeyer nexus is described from a single mature female 41.3 mm 
in SL, taken in 27-43 m from Ono-i-lau in the Lau Islands, Fiji. It can be distin- 
guished from other scorpaenids by the following combination of characters: 8 
short dorsal-fin spines, 3 spines and 8 segmented rays in the anal fin; 19 or 20 
pectoral-fin rays; | spine and 3 segmented rays in the pelvic fin; only unbranched 
rays in fins; no scales (except in lateral line); branchiostegal membranes not fused 
to isthmus; no preorbital spines; and no slit behind posteriormost hemibranch. 


A distinctive new genus and species of scorpaenid fish was collected from Ono- 
i-lau, Lau Islands, Fiji. The new form emphasizes the problems inherent in the 
most comprehensive and widely used classification of the scorpaenoids (Matsu- 
bara 1943a, b). It exhibits a number of advanced and primitive character-states 
not previously observed in combination in scorpaenoids, further obscuring the 
limits of several traditionally recognized but inadequately defined family-group 
taxa. The purpose of this paper is to make a name available for this species, 
deferring discussion of its relationships until the morphology of other presumably 
related scorpaenids can be more thoroughly studied. Assignment of this new 
species to the Scorpaenidae anticipates conclusions reached in studies now in 
progress by the first author. 

Methods of taking counts and measurements follow those of Eschmeyer (1969) 
as modified by Poss and Eschmeyer (1978). 

The right side of the holotype was dissected to reveal the configuration of the 
swimbladder musculature and to expose the dorsal bony elements of the first gill 
arch which were stained with alizarin but not removed. 


Eschmeyer, new genus 


Type-species.—Eschmeyer nexus. 

Diagnosis.—A scorpaenid fish with: dorsal fin VIII, 13; anal fin III, 8; pelvic 
fin I, 3; 19 or 20 pectoral-fin rays; extremely short anterior dorsal-fin spines; only 
unbranched rays in fins; frontal and parietal bones strongly ossified; no preorbital 
spines; no scales (except for lateral line); branchiostegal membranes not fused to 
isthmus; and no slit behind posteriormost hemibranch. 

Description.—Head large, slightly compressed (Figs. | and 2), with posterior 
half of frontal and parietal bones strongly ossified, rugose. Lacrimal (infraorbital 
1) immovable, with posteroventral border tapering to a small point. Nasal bone 
tubular, without spine. Mouth upturned. Teeth on vomer; none on palatines. No 
slit behind posteriormost hemibranch. Three infrapharyngobranchial tooth plates. 
Epibranchial of first gill arch with elongate uncinate process diverging from ramus 
of bone at angle of about 45° (Fig. 3; presence of interarcual cartilage uncertain). 
No cirri on lower jaw. Branchiostegal rays 7. Branchiostegal membranes of each 
side not fused to isthmus. Isthmus with fleshy extension posteriorly. Ventral 


310 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


surface of urohyal with broad transverse flange. Body compressed, oblong; skin 
slightly granular, scaleless, except for lateral line. Swimbladder absent. Swim- 
bladder musculature present, originating from cranium and cleithrum and insert- 
ing on parapophyses of vertebrae 6-10 (Fig. 4). Baudelot’s ligament attached to 
first vertebra. Neural spines of vertebrae 6—9 thin, almost hair-like (Fig. 5). Prox- 
imal dorsal-fin pterygiophores (except anteriormost 2) interdigitating without in- 
terruption between successive neural arches of precaudal vertebrae. Caudal skel- 
eton with parhypural and hypurals | and 2 fused into single autogenous plate, 
hypurals 3 and 4 fused and forming autogenous plate (slightly fused to urostyle?); 
hypural 5 autogenous; haemal spines of second and third preural centra autogen- 
ous; neural spine of second preural centrum short, 3 epurals; 2(1?) pairs of uro- 
neurals; hypurapophysis absent. 

Etymology.—Named for Dr. William N. Eschmeyer in recognition of his con- 
tributions to the study of scorpaenoid fishes; gender is masculine. 


Eschmeyer nexus, new species 
Figs. 1-5 


Holotype. —USNM 233855, 41.3 mm in standard length (SL). Fiji Is., Ono-i- 
lau I., outside of barrier reef on NW side of island 21°38’S, 178°45’W, 27—43 m, 
V.G. Springer party, Sta. VGS-82-14, | May 1982, 0705-1000 hrs. Mature female. 

Description.—Dorsal VIII, 13 (posteriormost ray split to base, counted as one); 
anal III, 8 (posteriormost split); pectoral 19 (left), 20 (right); pelvic I, 3; vertebrae 
I) +e ld = 2D. 

Anterior profile of head inclined about 39° from horizontal. Skin over lower 
part of head somewhat loose, wrinkled, and slightly granular in texture. Four 
infraorbital bones; second deep, with large subocular shelf on dorsomedial border 
(widest posteriorly). Third infraorbital bone deep, without spine, with shelf on 
dorsomedial border continuous with that on second infraorbital bone. Interorbit 
broad. Interorbital ridges weak, widely spaced, converging slightly but not meet- 
ing over midorbit. Supraorbital ridge vaguely defined, better ossified posteriorly. 
Anterior nostril prominent, pore situated before orbit, bordered anteroventrally 
by smaller pore of infraorbital sensory canal and anterodorsally by pore of su- 
praorbital canal. Laterosensory pores of head distinct, moderately sized, with 
slightly raised margins. Preopercle with 5 short, blunt spines, uppermost (first) 
most pungent, second largest, fifth a weak point. Pores of preopercular sensory 
canal with notably raised margins. Opercle strongly ossified, with 2 large, blunt 
spines on posterodorsal margin, smaller ventral spine more pungent; dorsal mar- 
gin inclined dorsoposteriorly about 5° above horizontal axis of body. Interopercle 
without spine. Parietal with broad low ridge. Pterotic with low ridge. Posttem- 
poral emarginate anteriorly, not forked; with strong spine. Supracleithrum bearing 
small, strong spine. Cleithral spine present. Ventral margin of dentary not strong- 
ly directed medially; no symphyseal knob. Mandibular pores 5 on each side, 
anteriormost 2 pores near symphysis arranged one behind the other. Anguloar- 
ticular bone not projecting strongly posteroventrally, inclined ventromedially at 
angle of about 45°. Maxilla without cirrus, extending to below middle of pupil. 
Gill rakers short, 14 total, 4 on upper arch, 10 on lower arch. Pseudobranch with 
11 filaments. 


VOLUME 96, NUMBER 2 311 


Fig. 1. Holotype of Eschmeyer nexus, USNM 233855, 41.3 mm SL. A, Lateral view; B, Dorsal 
view; C, Ventral view. 


312 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 2. Holotype of Eschmeyer nexus, USNM 233855. 


Lateral line not high on body, with 10 tubed scales, posteriormost on base of 
caudal fin. Lateral line scales without spinous projections. 

Dorsal fin originating over middle of operculum, spine in anterior part of fin 
extremely short, stout, and pungent. Dorsal-fin membrane weakly incised be- 
tween spines 3 and 4. Longest pectoral-fin rays (9-11 from dorsalmost) not reach- 
ing anus. Pelvic-fin insertion at base of pectoral fin. Pelvic-fin membrane adnate 
to body, fused to membrane of opposite side (see Comparisons below). Caudal 
fin stubby, rounded, with 18 total fin-ray elements (2 procurrent and 8 segmented 
dorsally, 7 segmented and 2 procurrent ventrally). 

Color in life unknown. Color pattern in 70% ethanol as in Fig. 1. Body color 
pale yellowish brown, covered by several broad dark-brown to brownish-black 
bars or patches. Two large saddle-shaped, somewhat irregular bars on upper two- 
thirds of body and dorsal-fin base; bars diffusely connected by narrower band 
just below lateral line; anterior bar below dorsal spines 5—8, posterior bar below 
segmented dorsal rays 2-9. Broad, dark greyish-brown patch on belly and lower 
part of pectoral-fin base, continuous with broad, nearly black, mostly submarginal 
band on lateral and medial surfaces of pectoral fins. Pale transverse band ex- 
tending across breast, just anterior to pectoral fins. Broad, uninterrupted brown- 
ish-black oblique band extending across caudal-fin base and distal part of seg- 
mented anal-fin rays. Narrower, but similarly colored, subterminal bar on caudal 
fin. Head with two brown bars radiating from eye; lower bar much broader, 
extending posteroventrally across cheek, over preopercle and lower half of oper- 
cle; upper bar narrower, directed dorsomedially and fused to slightly broader, 
small saddle-shaped bar extending transversely over dorsal-fin origin. Few irreg- 
ular dark-brown spots over anteroventral part of body and dorsal part of opercle. 
Few dark scattered specks in interorbit and over lower jaw. 


VOLUME 96, NUMBER 2 313 


i 


infrapharyngobranchial 


uncinate 
process 


epibranchial 


Fig. 3. Dorsolateral view of upper elements of right first gill arch in Eschmeyer nexus. Bar rep- 
resents | mm. 


Measurements for holotype (percent SL in parentheses): Head length 16.7(40); 
snout length 4.1(10); orbit diameter 3.9(9); interorbital width 3.2(8); upper jaw 
length 8.5(21); length of postorbital part of head 8.7(21); greatest body depth 
15.2(37); anal-fin length (base of first spine to end of longest ray) 13.9(34); caudal- 
fin length 9.0(22); pectoral-fin length 8.7(21); pelvic-fin length 7.3(18). Dorsal- 


EX 


PT 


Fig. 4. Dorsal (slightly oblique) view of Eschmeyer nexus swimbladder muscle and associated 
bony elements. Ribs, indicated by dashed line, lie dorsal to swimbladder muscle. BL, Baudelot’s 
ligament. CS, cranial slip of swimbladder muscle. EX, exoccipital. PA, vertebral parapophysis. PS, 
pectoral girdle slip of swimbladder muscle. PT, posttemporal. R,;, rib associated with 5 vertebral 
centrum. SO, spinooccipital nerve. V, ,9, 6, 10 vertebral centrum. Bar represents 1 mm. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


314 


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VOLUME 96, NUMBER 2 315 


spine lengths: first 1.5(4); second 1.9(5); third 3.2(8); fourth 3.9(9); fifth 3.5(8); 
penultimate 3.6(9); last 4.4(11). Anal-spine lengths: first 2.3(6); second 3.2(8); 
third 4.7(11). Distance between interorbital ridges 2.2(5); least depth of caudal 
peduncle 6.0(14). Distance from snout tip to: [base of] first dorsal spine (predorsal 
length) 15.1(37); second dorsal spine 16.2(39); third dorsal spine 17.5(42); fourth 
dorsal spine 19.9(48); fifth dorsal spine 21.8(53); pelvic-fin insertion 14.2(34). 
Transverse width of first dorsal spine at midlength 0.2(0.5); deepest incision of 
fin membrane between third and fourth dorsal spines (from tip of fourth spine to 
membrane) 1.0(2); length of dorsalmost preopercular spine 1.4(3). Distance be- 
tween: tip of opercle and dorsal-fin base 5.2(13); first and fifth dorsal spines 
6.1(15); fifth and last dorsal spines 4.8(12); last dorsal spine and last dorsal ray 
6.2(15); anal-fin origin and last anal ray 10.7(26); pelvic-fin insertion and anal-fin 
origin 16.8(41); first dorsal spine and pelvic-fin insertion 14.0(34); first dorsal spine 
and anal-fin origin 21.2(51); fifth dorsal spine and pelvic-fin insertion 15.9(38); last 
dorsal spine and pelvic-fin insertion 17.5(42); last dorsal spine and last anal ray 
19.2(46); last dorsal ray and anal-fin origin 13.7(33); last dorsal spine and anal-fin 
origin 13.8(33). 

Etymology.—From the Latin nectere (to tie or connect) in reference to features 
of the species that, in combination, suggest a close relationship to several groups 
of scorpaenoids. Here treated as a noun in apposition. 

Remarks.—The holotype was captured by use of SCUBA and rotenone in an 
area usually having strong wave action. Especially calm weather made diving at 
this locality possible. Rotenone was placed in a large gently downsloping channel 
in the reef slope at a depth of about 27 m and on the steep sloping face of the 
reef at about 36 m. Dead fishes of a number of species were picked up to a depth 
of about 43 m, but most were obtained from the channel at the shallowest depth. 

Comparisons.—Eschmeyer can be differentiated from all other scorpaenid fish- 
es by the combination of characters given in the diagnosis. Among scorpaenids 
only Minous Cuvier (Scorpaenidae: Minoinae) has as few as 8 dorsal spines (7— 
12). Eschmeyer differs conspicuously from Minous in having 3 rather than 5 
segmented rays in the pelvic fin and in lacking a detached ray in the pectoral fin, 
branchiostegal membranes that are fused to the isthmus, preorbital spines, and a 
movable lacrimal. Among scorpaenoids, Eschmeyer is perhaps most similar to 
Peristrominous Whitley (Aploactinidae) in overall appearance but can be distin- 
guished readily in having 8 dorsal-fin spines (12 or 13 in Peristrominous), 19 or 
20 pectoral-fin rays (14 or 15 in Peristrominous), 3 anal-fin spines (1 in Peris- 
trominous), and in lacking a fleshy extension at the anteriormost part of the 
isthmus. 

Among other scorpaenids, fusion of the pelvic-fin membranes to each other is 
an atypical condition known to occur only rarely in some specimens of Gymnap- 
istes marmoratus (Cuvier) and Vespicula dracaena (Cuvier), both currently as- 
signed to the Tetraroginae. Whether the holotype of Eschmeyer nexus is such a 
variant is unknown. 


Acknowledgments 


Funds for support of the fieldwork resulting in the collection of Eschmeyer 
nexus and publication of this study were derived from grants made to V. G. 
Springer by the Smithsonian Scholarly Studies Program and the Max and Victoria 


316 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Dreyfus Foundation. We thank Dr. William Smith-Vaniz for reviewing this manu- 
script. 
Literature Cited 


Eschmeyer, W. N. 1969. A systematic review of the scorpionfishes of the Atlantic ocean (Pisces: 
Scorpaenidae).—Occasional Papers of the California Academy of Sciences 79:1—143, figs. I- 


13. 
Matsubara, K. 1943a. Studies on the scorpaenoid fishes of Japan. Anatomy, phylogeny and tax- 
onomy (1I).—Transactions of the Sigenkagaku Kenkyusyo 1:1—170, figs. 1-66. 


—. 1943b. Studies on the scorpaenoid fishes of Japan. Anatomy, phylogeny and taxonomy 
(I1I1).—Transactions of the Sigenkagaku Kenkyusyo 2:171—486, figs. 67-156, 4 plates. 

Poss, S. G., and W. N. Eschmeyer. 1978. Two new Australian velvetfishes, genus Paraploactis 
(Scorpaeniformes: Aploactinidae), with a revision of the genus and comments on the genera 
and species of the Aploactinidae.—Proceedings of the California Academy of Sciences 41(18): 
401-426, fig. 1-14, 6 tbs. 


(SGP) Department of Ichthyology, The Academy of Natural Sciences of Phila- 
delphia, Nineteenth and the Parkway, Philadelphia, Pennsylvania 19103. (VGS) 
Division of Fishes, National Museum of Natural History, Washington, D.C. 20560. 


PROC. BIOL. SOC. WASH. 
96(2), 1983, pp. 317-322 


FENNEROSQUILLA, A NEW GENUS OF STOMATOPOD 
CRUSTACEAN FROM THE NORTHWESTERN ATLANTIC 


Raymond B. Manning and David K. Camp 


Abstract.—A new genus, Fennerosquilla, is recognized for Chloridella hep- 
tacantha Chace, 1939, a species previously assigned to the genus Squilla. This 
new genus can be distinguished from Squilla by the presence of seven teeth on 
the dactylus of the claw and by the strongly bilobed lateral process of the sixth 
thoracic somite. 


The unusual morphological features of the northwestern Atlantic Chloridella 
heptacantha Chace (1939:52), subsequently assigned to the genus Squilla Fabri- 
cius, 1787, have been recognized for some time (Manning 1969:147; 1978b:40). It 
is the only species now assigned to Squilla with seven teeth on the dactylus of 
the claw and a strongly bilobed lateral process of the sixth thoracic somite. Recent 
collections of this species (Camp 1983) led us to reevaluate its generic status. We 
believe that it is sufficiently distinct from other species assigned to Squilla to 
warrant the recognition of a new, monotypic genus. Descriptors used to indicate 
body size and eye size follow definitions given by Manning (1978b:2). 


Fennerosquilla, new genus 
Fig. 1 


Definition.—Size moderate, total lengths of adults less than 100 mm. Body 
generally smooth, surface slightly punctate; carinae well developed, sides eroded. 
Anterior margin of ophthalmic somite rounded or faintly emarginate, unarmed. 
Eye small to very small, cornea bilobed, set obliquely on stalk. Ocular scales 
subquadrate to emarginate, separate. Rostral plate elongate, triangular, with me- 
dian carina. Carapace with normal complement of carinae (median, intermediates, 
laterals, reflected marginals), median with well-developed anterior bifurcation. 
Mandibular palp present. Four epipods present. Dactylus of claw with 7 teeth. 
Lateral process of fifth thoracic somite a single broad lobe, directed laterally. 
Lateral process of sixth thoracic somite strongly bilobed, subtriangular anterior 
lobe almost as large as posterior. Lateral process of seventh thoracic somite 
bilobed, triangular anterior lobe smaller than posterior. Abdomen with 8 carinae 
(paired submedians, intermediates, laterals, and marginals) on each of anterior 5 
somites, intermediates and laterals strongly developed. Telson inflated basally, 
dorsal surface with median carina and carinae of marginal teeth, lacking supple- 
mentary dorsal carinae or tubercles; 3 pairs of marginal teeth present, submedians 
with fixed apices; prelateral lobes present. Basal prolongation of uropod produced 
into 2 strong spines, inner longer; inner margin crenulate, unarmed. 

Type-species.—Chloridella heptacantha Chace, 1939, herein designated. The 
genus is monotypic. 

Etymology.—We believe it is appropriate to dedicate this genus to Fenner A. 
Chace, Jr., who originally described the type-species. The gender is feminine. 


318 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 1. Fennerosquilla heptacantha, female, total length 74.5 mm, USNM 126025. a, Head and 
anterior part of carapace; b, Exposed thoracic somites 5-7; c, Carpus of raptorial claw; d, Propodus 
and dactylus of raptorial claw; e, Sixth abdominal somite, telson and right uropod, dorsal view; f, 
Telson, right lateral view; g, Right uropod, ventral view. 


Remarks.—Until now, 25 genera had been recognized in the family Squillidae 
(see Manning 1980:367 for a list of 24 of these genera and original references, and 
Manning and Lewinsohn 1982:352), most having been separated from the genus 
Squilla in the last two decades (Manning 1968, 1972, 1976, 1977, 1978a, b). Of 
the 26 genera now assigned to the Squillidae, eleven are distinguished by the 
presence of movable apices on the submedian teeth of the telson in adults; in the 
remaining 15 genera (including Fennerosquilla), the apices of the submedian teeth 
are fixed in adults, although they may be movable in postlarvae and early juvenile 


VOLUME 96, NUMBER 2 319 


stages. Of these latter 15 genera, only five have single lateral processes on the 
fifth thoracic somite: Anchisquilla, Lenisquilla, Squilloides, Squilla, and Fen- 
nersoquilla. The first two of these genera completely lack the median carina of 
the carapace. Squilloides lacks a mandibular palp and has but four teeth on the 
dactylus of the claw. Thus, Fennerosquilla might be confused only with members 
of the genus Squilla. 

Fennerosquilla can be readily distinguished from Squilla by the following dis- 
tinctive features: elongate, triangular rostral plate; seven teeth on the claw; re- 
duction of number of epipods to four; and large anterior lobe of the sixth thoracic 
somite. Other features characteristic of Fennerosquilla heptacantha serving to 
distinguish it from Squilla include: long, slender antennular peduncles, longer 
than the carapace and rostral plate combined; extremely narrow, elongate cara- 
pace, with the anterior width less than half the median length; long, slender 
propodus of the raptorial claw (the claw, when folded, exceeds the anterior mar- 
gin of the ophthalmic somite); strongly flattened, strongly carinate abdomen; 
basally inflated telson, with the median carina appearing convex anteriorly, con- 
cave posteriorly in lateral view; and relatively large rounded lobe on the outer 
margin of the inner spine of the basal prolongation of the uropod. Finally, the 
eyes are small relative to body size (Table 1); eyes of the smallest F. heptacantha 
are similar in relative size to those of the largest specimens of Squilla (except 
possibly §. surinamica, a much smaller species than F. heptacantha and most 
other species of Squilla). 

We suspect that the presence of a strongly bilobed lateral process of the sixth 
thoracic somite in F. heptacantha is not an indication of affinity with Indo-West 
Pacific stocks of squillids, most of which share this feature, but is an independent 
development. Fennerosquilla may represent an offshoot of the genus Squilla. 

The importance of shape of lateral processes of the exposed thoracic somites 
in distinguishing groups of species within the Squillidae has been recognized since 
Kemp (1913:20, 21) divided the long-recognized, heterogeneous Squilla into two 
broad groups based solely on shape of the lateral process of the fifth thoracic 
somite. In a later revision of the Squillidae (see Manning 1968), the nature of 
differences in shape of the lateral process of the fifth thoracic somite among 
various squillid species was thoroughly discussed. At that time, two basic forms 
of the lateral process were recognized, and the difference between them was 
believed to indicate a fundamental difference between two stocks within the fam- 
ily. One form was expressed as a bilobed process of the fifth thoracic somite; the 
two lobes were either on the same horizontal plane, as in Alima, or almost so, 
and consequently there was no ventral spine on that somite. In the second form, 
the anterior lobe had rotated ventrally in relation to the posterior lobe, was no 
longer basally connected with the posterior lobe, and formed a separate ventral 
spine. Thus, the posterior lobe projected laterally or anterolaterally from the 
body, but the anterior lobe projected more ventrally and often was obscured from 
dorsal view because of its position on the body. Based on this fundamental dif- 
ference, as well as other considerations, several new genera were erected to 
contain species formerly assigned to Squilla and having a bilobed lateral process 
of the fifth thoracic somite. Species retained in Squilla had a ventral spine and a 
single-lobed lateral process of the fifth thoracic somite. One of the genera then 
recognized, Oratosquilla Manning, 1968, received the greatest number of species 


320 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Table |.—Representative corneal indices (CI = carapace length divided by cornea width, multiplied 
by 100) of several species of Squilla compared with those of F. heptacantha. 


Species Known range of CI 
Squilla lijdingi Holthuis, 1959 260-363 
S. rugosa Bigelow, 1893 270-424 
S. empusa Say, 1818 296-491 
S. deceptrix Manning, 1969 311-446 
S. caribaea Manning, 1969 315-405 
S. intermedia Bigelow, 1893 316-400 
S. chydaea Manning, 1962 328-413 
S. edentata edentata (Lunz, 1937) 338492 
S. obtusa Holthuis, 1959 350-421 
S. cadenati Manning, 1970 352-468 
S. brasiliensis Calman, 1917 354-492 
5S. discors Manning, 1962 361-439 
S. neglecta Gibbes, 1850 369-498 
S. edentata australis Manning, 1969 370-447 
S. mantis (Linnaeus, 1758) 38 1-433 
Fennerosquilla heptacantha (Chace, 1939) 412-576 
S. surinamica Holthuis, 1959 441-469 


having a bilobed process. Species of Squilla sensu stricto were then restricted to 
the Atlanto-East Pacific, whereas species of Oratosquilla were confined almost 
exclusively to the Indo-West Pacific region [except O. massavensis (Kossmann, 
1880), an immigrant to the eastern Mediterranean Sea through the Suez Canal 
(Lewinsohn and Manning 1980:11)]. 

Differences in shape of the lateral processes of the sixth and seventh thoracic 
somites were not stressed in the 1968 revision of the Squillidae, although it was 
suggested that those features “‘may prove to be of some importance in classifi- 
cation at the generic level’’ (Manning 1968:118). All of the species transferred to 
Oratosquilla in 1968 share a strongly bilobed lateral process of the sixth thoracic 
somite, whereas only one of the species then retained in Squilla, S. heptacantha, 
has a strongly bilobed process on that somite (Manning 1969: fig. 47d; Fig. 1b, 
herein). In that species, the anterior lobe of the process of the sixth thoracic 
somite is almost as large as the posterior lobe. In other species retained in Squilla, 
the lateral process of the sixth thoracic somite is either a single lobe, or the 
anterior lobe is distinctly smaller than the posterior one. Further, most species 
transferred to Oratosquilla in 1968 have a bilobed lateral process of the seventh 
thoracic somite as well, and the anterior lobe of that process is relatively large. 
Species retained in Squilla have either a simple process of the seventh thoracic 
somite or a bilobed process with the anterior lobe greatly reduced. 

After the 1968 revision of the Squillidae and a subsequent monograph of the 
West Atlantic species were completed (Manning 1969), more detailed studies were 
begun on species of Oratosquilla, and the genus was found to comprise several 
morphologically heterogeneous groups of species (Manning 1971). As more spec- 
imens became available and additional information on their morphology was ob- 
tained, several new genera were recognized for species previously assigned to 
Oratosquilla. In some of these genera, basic distinctions from Oratosquilla in- 


VOLUME 96, NUMBER 2 32] 


cluded different configurations of the lateral processes of the exposed thoracic 
somites and different numbers of teeth on the dactylus of the claw. For example, 
Busquilla Manning, 1978a, was erected for O. quadraticauda (Fukuda, 1911) (not 
seen by Manning in 1968) and another, previously undescribed species. The lat- 
eral process of the fifth thoracic somite in these species, although appearing in 
dorsal view to be bilobed with both lobes in almost the same plane, is actually 
somewhat similar to the lateral process found in species of Squilla; the two lobes 
are separate and in different planes as well. Furthermore, in Busquilla species 
the anterior lobes of the lateral processes of the sixth and seventh somites are 
proportionally smaller than those seen in most species of Oratosquilla, and their 
claws have five rather than six teeth on the dactylus. 

Later, Kempina Manning, 1978b, was erected for O. mikado (Kemp and Cho- 
pra, 1921) and two other species having lateral processes with separated lobes in 
different planes on the fifth thoracic somite and strongly bilobed processes on 
the sixth and seventh somites. Like species of Oratosquilla, species of Kempina 
have six teeth on the dactylus of the claw. Natosquilla Manning, 1978b, erected 
for O. investigatoris (Lloyd, 1907), also has separate lobes in different planes on 
the fifth somite, but the anterior lobes of the lateral processes of the following 
two somites are reduced in size. Natosquilla further differs from Oratosquilla in 
having 10-18 rather than six teeth on the dactylus of the claw. 

These findings have clarified the usefulness of shapes of lateral thoracic pro- 
cesses as generic characters. First, the concept that two distinct stocks within 
the Squillidae can be distinguished by the shape of the lateral process of the fifth 
thoracic somite is still valid but requires modification. The principally Indo-West 
Pacific stock, plus the wide-ranging Alima, can no longer be characterized as 
having both lobes of the lateral process of the fifth thoracic somite basally con- 
nected and in the same or almost the same horizontal plane. However, this stock 
can be differentiated from the principally Atlanto-East Pacific stock, comprising 
Squilla and its allies, by whether or not the two lobes are visible in dorsal view. 
In the latter group, the posterior lobe always obscures the ventrally rotated an- 
terior lobe, whereas in the former group, even though the anterior lobe may be 
ventrally rotated, it is still visible in dorsal view. Second, as demonstrated by 
Busquilla and Fennerosquilla, shape of the lateral process of the sixth thoracic 
somite may be an important generic character (in combination with other char- 
acters), as suggested in 1968. Finally, shape of the process of the seventh somite 
alone is not necessarily important at the generic level, as shown by some species 
of Oratosquilla. In O. asiatica Manning, 1978, O. pentadactyla Manning, 1978, 
O. quinquedentata (Brooks, 1886), and O. woodmasoni (Kemp, 1911), the an- 
terior lobe of the lateral process of the seventh thoracic somite is proportionally 
much smaller than in other species of the genus. 

Since 1968, when Squilla was restricted to its Atlanto-East Pacific components, 
several other features have emerged as important characters at the generic level 
within the Squillidae, including number of epipods, presence or absence of the 
mandibular palp, and number of teeth on the dactylus of the claw. As a general 
rule, these features are constant at the generic level. One possible exception 
occurs among species assigned to Clorida, where, so far as is known, all of these 
features vary. That genus, however, also requires revision. 

We plan to reevaluate these characters in species now assigned to Squilla, in 


322 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


which two species (S. neglecta Gibbes, 1850, and S. parva Bigelow, 1891) lack 
the mandibular palp, three species ($. aculeata Bigelow, 1893, S. prasinolineata 
Dana, 1852, and §. grenadensis Manning, 1969) have four rather than five epi- 
pods, and another species (S. decimdentata Manning, 1970) has 10 rather than 
six teeth on the claw. Two of these species, $. neglecta and S. prasinolineata, 
have but five teeth on the claw. 


Acknowledgments 


We thank Lilly King Manning for preparing the illustrations, and W. G. Lyons, 
J. F. Quinn, Jr., and E. W. Truby of the Florida Department of Natural Resources, 
Marine Research Laboratory, for their comments on the manuscript. 


Literature Cited 


Camp, D. K. 1983. Occurrence of Squilla heptacantha (Chace, 1939) (Crustacea: Stomatopoda: 
Squillidae) in the northeastern Gulf of Mexico.—Northeast Gulf Science 6(1):55—57. 

Chace, F. A., Jr. 1939. Preliminary descriptions of one new genus and seventeen new species of 
decapod and stomatopod Crustacea. Reports on the scientific results of the first Atlantis ex- 
pedition to the West Indies, under the joint auspices of the University of Havana and Harvard 
University.—Memonias de la Sociedad Cubana de Historia Natural 13(1):31—54. 

Kemp, S. 1913. An account of the Crustacea Stomatopoda of the Indo-Pacific region, based on the 
collection in the Indian Museum.—Memoirs of the Indian Museum 4: 1-217. 

Lewinsohn, Ch., and R. B. Manning. 1980. Stomatopod Crustacea from the eastern Mediterra- 
nean.—Smithsonian Contributions to Zoology 305: 1-22. 

Manning, R. B. 1968. A revision of the family Squillidae (Crustacea, Stomatopoda), with the de- 
scription of eight new genera.—Bulletin of Marine Science 18(1): 105-142. 

——. 1969. Stomatopod Crustacea of the western Atlantic.—Studies in Tropical Oceanography 
8:vill + 380 pp. 

——. 1971. Keys to the species of Oratosquilla (Crustacea: Stomatopoda), with descriptions of 
two new species.—Smithsonian Contributions to Zoology 71:1—16. 

——. 1972. Notes on some stomatopod crustaceans from Peru.—Proceedings of the Biological 
Society of Washington 85:297—307. 

—. 1976. Redescriptions of Oratosquilla indica (Hansen) and Clorida verrucosa (Hansen), with 
accounts of a new genus and two new species (Crustacea, Stomatopoda).— Beaufortia 25(318): 
1-13. 

—.. 1977. Preliminary accounts of five new genera of stomatopod crustaceans.—Proceedings 
of the Biological Society of Washington 90:420-423. 

——. 1978a. New and rare stomatopod Crustacea from the Indo-West-Pacific region.—Smith- 
sonian Contributions to Zoology 264: 1—36. 

——. 1978b. Further observations on Oratosquilla, with accounts of two new genera and nine 
new species (Crustacea: Stomatopoda: Squillidae)—Smithsonian Contributions to Zoology 
272: 1-44. 

——. 1980. The superfamilies, families, and genera of Recent stomatopod Crustacea, with diag- 

noses of six new families.—Proceedings of the Biological Society of Washington 93:362-372. 

, and Ch. Lewinsohn. 1982. Rissoides, anew genus of stomatopod crustacean from the East 

Atlantic and South Africa.—Proceedings of the Biological Society of Washington 95:352-353. 


(RBM) Department of Invertebrate Zoology, National Museum of Natural His- 
tory, Smithsonian Institution, Washington, D.C. 20560; (DKC) Florida Depart- 
ment of Natural Resources, Marine Research Laboratory, 100 Eighth Avenue 
S.E., St. Petersburg, Florida 33701. 


PROC. BIOL. SOC. WASH. 
96(2), 1983, pp. 323-332 


PROCAMBARUS (ORTMANNICUS) LEITHEUSERI, NEW 
SPECIES, ANOTHER TROGLOBITIC CRAYFISH 
(DECAPODA: CAMBARIDAE) FROM 
PENINSULAR FLORIDA 


Richard Franz and Horton H. Hobbs, Jr. 


Abstract.—A new troglobitic crayfish, Procambarus (Ortmannicus) leitheuseri, 
is described from six localities in Hernando and Pasco counties, Florida. Its 
closest affinities seem to be with the two subspecies of P. (O.) lucifugus. It may 
be distinguished from the nominate subspecies by the presence of pigment in the 
eye, and from P. (O.) |. alachua by features of the first pleopod of the male, 
among which is the absence of a caudal process. 


Twelve troglobitic crayfishes belonging to three genera are now known to in- 
habit the subterranean waters of Florida. Except for Procambarus (Ortmannicus) 
franzi Hobbs and Lee (1976) and the new species described herein, all were in- 
cluded in a review of the American troglobitic decapods (Hobbs et al., 1977). These 
two crayfishes not reported there occur in the peninsular section of the state: P. 
(O.) franzi, known only from the type-locality, Orange Lake Cave (Sec. 33/34, 
T. 12S, R. 21E) and Hell Hole (Sec. 6, T. 14S, R. 21E), Marion County, and P. 
(O.) leitheuseri, new species, from six localities in Hernando and Pasco counties. 
The latter records mark the southwestern limits of the known range of the trog- 
lobitic members of the subgenus Ortmannicus in peninsular Florida. A key to the 
albinistic crayfishes of the subgenus is appended to the description that follows. 

We are pleased to name this crayfish for its discoverer Arthur T. Leitheuser, 
who has added much to our knowledge of the distribution of the troglobitic cray- 
fishes of Florida. 


Procambarus (Ortmannicus) leitheuseri, new species 
Fig. | 


Diagnosis.—Albinistic, eyes without facets but provided with small pigment 
spot. Rostrum with marginal spines; median carina absent. Carapace with cervical 
spine cephaloventral to row of small spines or tubercles flanking caudal margin 
of cervical groove. Aerola 6.3 to 10.4 times as long as broad and constituting 32.9 
to 38.2% of total length of carapace (43.8 to 49.7% of postorbital carapace length). 
Suborbital angle absent. Postorbital ridge with cephalic spine and with or without 
2 to several more posterior spines or tubercles. Hepatic area with many small 
tubercles, some spiniform. Antennal scale about twice as long as wide, broadest 
Slightly distal to midlength. Ischia of third and fourth pereiopods of first form 
male with simple hooks, that on third overreaching basioischial articulation and 
that on fourth highly arched, almost reaching basioischial articulation but lacking 
opposing tubercle on basis; coxa of fourth pereiopod with prominent oblique boss. 
First pleopod of first form male reaching coxa of third pereiopod, asymmetrical, 
provided with subapical setae; distal extremity bearing subspiculiform mesial 


324 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


process directed caudally at approximately 75 degree angle to shaft of appendage 
and curved somewhat laterally; cephalic process acute, somewhat hooding central 
projection cephalically, and directed caudodistally; caudal element lacking caudal 
knob but represented by prominent, corneous adventitious process caudome- 
sially, latter rounded distally, convex mesially, and somewhat concave laterally; 
and corneous beaklike central projection, most conspicuous of terminal elements, 
directed caudodistally subparallel to cephalic process. Annulus ventralis freely 
movable, subrhomboidal, about twice as broad as long, and completely exposed, 
not partly hidden by projections from sternum immediately cephalic to it; cephalic 
area with convex, elevated marginal area bearing submedian, oblique furrow; 
sinus originating in furrow and following sigmoid curve, terminating almost on 
median line slightly posterior to midlength of annulus. Postannular sclerite slightly 
more than half as wide and about half as long as annulus with cephalomedian 
area somewhat inflated. First pleopod in female moderately well developed. 

Holotypic male, form 1.—Cephalothorax (Fig. la, k) subcylindrical. Abdomen 
narrower than thorax (8.4 and 10.0 mm). Greatest width of carapace greater than 
height at caudodorsal margin of cervical groove. Areola 8.4 times as long as wide 
with | or 2 punctations across narrowest part. Cephalic section of carapace ap- 
proximately 1.6 times as long as areola, length of latter 38.2% of entire length of 
carapace (48.0% of postorbital carapace length). Rostrum with margins subpar- 
allel along caudal half and gently convergent to base of acumen where provided 
with small slightly divergent marginal spines. Acumen reaching base of distal 
fourth of ultimate segment of antennular peduncle; dorsal surface excavate and 
punctate. Subrostral ridge weak and evident in dorsal aspect along caudal seventh 
of rostrum. Postorbital ridges well developed, grooved dorsolaterally, and ter- 
minating cephalically in small corneous spine; posteriormost part of right ridge 
with 3 minute corneous subacute tubercles, left with 5, two of which somewhat 
lateral to axis of ridge. Dorsolateral part of cervical groove flanked caudally by 
row of spiniform tubercles, ventralmost member of row (cervical spine) larger 
than others. Suborbital angle obsolete. Branchiostegal spine moderately strong 
and acute. Except for anterior parts of orbital, antennal, and mandibular areas, 
entire dorsolateral and lateral surfaces of carapace studded with tubercles, some 
of which spiniform. 

Abdomen slightly longer than carapace (23.5 and 22.0 mm). Pleura of third 
through fifth abdominal segments rounded anteroventrally and angular caudo- 
ventrally. Cephalic section of telson with 2 spines in each caudolateral corner, 
more mesial one in each movable. Cephalic lobe of epistome (Fig. 17) cordiform, 
with cephalolateral margins slightly elevated (ventrally); main body with ante- 
riorly flared depression but lacking distinct fovea; epistomal zygoma broadly 
arched. Ventral surface of proximal podomere of antennular peduncle with spine 
slightly distal to midlength. Antenna with prominent spine on lateral surface of 
basis and another on ventral surface of ischium; flagellum extending caudally 
beyond telson by almost 4 times length of latter. Antennal scale (Fig. 17) about 
twice as long as broad, widest slightly distal to midlength, and lamellar area about 
3 times as wide as thickened lateral part. 

Third maxilliped overreaching rostrum by length of ultimate podomere; ischium 
with distolateral margin subserrate, ending in acute prominence, and lateral half 


VOLUME 96, NUMBER 2 325 


Fig. 1. Procambarus (O.) leitheuseri (all illustrations from holotype except d, f, from morphotype, 
and e from allotype): a, Lateral view of carapace; b, d, Mesial view of first pleopod; c, Caudolateral 
view of first pleopod; e, Annulus ventralis; f-h, Lateral view of first pleopod; i, Antennal scale; j, 
Epistome; k, Dorsal view of carapace; /, Proximal podomeres of third, fourth, and fifth pereiopods; 
m, Dorsal view of distal podomeres of cheliped; n, Caudal view of first pleopods. 


of ventral surface with scattered short setiferous punctations; exopod reaching 
almost to end of merus. 

Right chela (Fig. 1m) subovate in cross section, not strongly depressed. Mesial 
surface of palm with several irregular rows of 12 to 14 strongly elevated tubercles; 


326 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


remainder of palm strongly tuberculate. Both fingers with well defined submedian 
longitudinal ridge flanked proximally by tubercles and more distally by setiferous 
punctations. Opposable margin of fixed finger with row of 9 tubercles along prox- 
imal half of finger, fourth from base largest, and with single, large, more ventrally 
situated one slightly proximal to midlength; row of minute denticles extending 
between and ventral to row of tubercles from base of finger almost to corneous 
tip. Lateral surface of fixed finger with tubercles, decreasing in size distally, along 
proximal half followed by row of setiferous punctations, latter reaching corneous 
tip of finger. Opposable margin of dactyl with row of 11 corneous tubercles, sixth 
from base largest, along proximal two-fifths of finger; minute denticles between 
tubercles and forming broad band immediately distal to last tubercle of row, band 
continuing to base of corneous tip of finger. Mesial surface of dactyl similar to 
lateral surface of fixed finger; tubercles, diminishing in size distally, reaching 
almost midlength of finger. 

Carpus of cheliped longer than broad, tuberculate on all surfaces although 
sparsely so ventrally; 3 most prominent tubercles spiniform, one situated on 
dorsodistal angle and 2 on ventrodistal margin. Shallow oblique sulcus on dorsal 
surface flanked by small subsquamous tubercles. 

Merus of cheliped strongly tuberculate except for proximal parts of mesial and 
lateral surfaces. Dorsal surface with sublinear series of tubercles; those on ventral 
surface not limited to usual 2 rows; rows poorly defined but mesial one consisting 
of about 25, more distal members of lateral row larger and corneous. Ischium 
with irregular row of 5 tubercles along mesial margin. 

Hooks on ischia of third and fourth pereiopods (Fig. 1/) as described in Di- 
agnosis. Coxa of fourth pereiopod with prominent oblique (almost vertically 
disposed), somewhat inflated boss, its ventral border turned mesially; boss on 
coxa of fifth pereiopod much smaller and tuberculiform. 

Sternum between third and fourth pereiopods rather deep with conspicuous 
mat of plumose setae extending mesially from ventrolateral margins. First pleo- 
pods (Fig. 1b, c, g, h, n) as described in Diagnosis. Uropods with both lobes 
of basal podomere bearing spines, that on mesial lobe very strong; distomedian 
spine on mesial ramus far removed from distal margin of ramus. 

Allotypic female.—Differing from holotype in following respects: areola 9 times 
as long as wide; cephalic section of carapace 1.8 times length of areola, latter 
only 35.2% of entire length of carapace, 45.3% of postorbital carapace length; 
acumen with accessory spine on dextral side at base of corneous tip, latter slightly 
overreaching peduncle of antennule; posterior tubercles of postorbital ridges weak 
and none corneous; posteroventral extremities of third and fourth abdominal 
segments not distinctly angular; epistome with paired marginal angles flanking 
anteromedian projection; antennal flagellum slightly shorter than in holotype; 
serrations on lateral margin of ischium of third maxilliped less well developed 
than in holotype; opposable margin of fixed finger of chela with row of 15 tuber- 
cles along proximal fourth, third from base largest; tubercles on lateral margin of 
finger very poorly developed; opposable margin of dactyl with row of 8 tubercles, 
fifth from base largest; irregular mesial and lateral rows of tubercles on ventral 
surface of merus consisting of about 16 and 18, respectively. 

Annulus ventralis (Fig. le) only moderately deeply located on sternum (see 


VOLUME 96, NUMBER 2 327 


Diagnosis). First pleopod reaching cephalic margin of annulus when abdomen 
flexed. (See Measurements.) 

Morphotypic male, form IT.—Differing from the holotype, except in secondary 
sexual characters, in only few respects: spines and tubercles almost everywhere 
decidedly more prominent, but little different in number or distribution; pigment 
spot in eye larger and pigment more concentrated; acumen overreaching anten- 
nular peduncle by about one-third of its length; subrostral ridges evident dorsally 
along almost basal half of rostrum; 4 well developed spines along posterior part 
of postorbital ridge, left with 3; hooks on ischia of third and fourth pereiopods 
much less well developed, and bosses on coxae of fourth and fifth pereiopods 
not nearly so prominent. 

First pleopods (Fig. Id, f) only slightly asymmetrical and with distinct hump 
on preapical cephalic surface. Mesial and cephalic processes strong, and central 
projection less prominent than in first form male; all disposed much as in holo- 
type, but more distinctly caudally; caudal element not clearly defined. 

Type-locality.—‘*Eagle’s Nest’? (= Lost Sink, Eagle Hole), 5.4 km northwest 
of the junction of U.S. Highway 19 and State Road 50 (NE4, NE%4, SW", Sec. 
21, T. 22S, R. 17E), Hernando County, Florida. This is a flooded cave system 
that opens to the surface in the bottom of a sinkhole pond 76 meters in diameter, 
in the Chassahowitzka Swamp. The entrance consists of a series of vertical shafts, 
most of which are either too small to permit access or are blocked by rocks. The 
largest shaft, 2 by 2 meters in diameter, opens into the top of a very large bell- 
shaped gallery at a water depth of 21 meters. The room is roughly oval in shape 
with the long axis oriented in an east-west direction. Tunnels lead out on the 
east and west sides, and water flow is from east to west, although some divers 
have reported observing occasional reversals in this pattern. The highest part of 
the room’s floor is at a water depth of 38 meters, but the floor quickly drops away 
on all sides to depths of as much as 61 meters. The floors of the room and tunnels 
are composed of coarse sand and, in places, of large blocks of breakdown. Easily 
disturbed organic silt, sometimes in thick layers, covers the floor under the en- 
trance shafts and in less traveled, deeper sections of the cave. It is present in 
cracks and crevices in the ceiling of the room. Divers have explored both the 
upstream and downstream passages to depths in excess of 91 meters and distances 
in excess of 300 meters. 

Disposition of types.—The holotype, allotype, and morphotype are deposited 
in the National Museum of Natural History (Smithsonian Institution), numbers 
178361, 178362, and 178585, respectively, as are paratypes consisting of two 
second form males (maintained alive) and a dry female. The remaining paratypes 
(see specimens examined) are deposited in the Florida State Museum. 

Size.—The largest of the available specimens is a female from Die Polder 3 which 
has a carapace length of 27.4 (postorbital carapace length, 20.8) mm. Correspond- 
ing lengths of the largest male (form II, from Arch Sink) are 24.4 and 18.6 mm. 
The holotype is the only first form male available (see Tab. | for measurements). 

Range and specimens examined.—Procambarus (O.) leitheuseri has been col- 
lected in only six localities, all in Hernando and Pasco counties, Florida: HER- 
NANDO COUNTY—(1) the type-locality, 1 ¢I, the holotype, A. T. Leitheuser 
and L. F. Collins, coll.; (2) Die Polder 2, 3.6 km ENE of junction of U.S. Hwy. 


328 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Table 1—Measurements (mm) of Procambarus (O.) leitheuseri. 
Holotype Allotype Morphotype 

Carapace 

Height 8.9 V3 10.0 

Width 10.0 8.0 11.1 

Total length 22.0 17.9 24.4 

Postorbital length 17.5 13.9 18.6 
Areola 

Width 1.0 0.7 1.2 

Length 8.4 6.3 8.8 
Rostrum 

Width B)a72 2.4 Bis), 

Length del) 4.9 6.6 
Right Chela 

Length, palm mesial margin Ted 4.2 6.1 

Palm width 3.7 2.4 3.6 

Length, lateral margin Ol 12.8 17.9 

Dactyl length 12.8 Tes) 10.8 
Abdomen 

Width 8.4 6.7 9.3 

Length 235) 19.1 24.8 


19 and State Rte. 50 (Weekiwachee Springs) (NW'4, NE'4, NE", Sec. 5, T. 22S, 
R. 18E), depth about 60 m, near bottom of shaft, 1 2, 24 February 1980, A. T. 
Leitheuser and J. Bentz, coll.; (3) Die Polder 3, 4.2 km E of junction of U.S. 
Hwy. 19 and State Rte. 50 (Weekiwachee Springs) (NW'4, SE’44, NW, Sec. 5, 
T.23S, R.18E), 1 2, 5 September 1982, W. K. Fehring, S. Fehring, coll.; 4 2, 9 
October 1982, WKF, SF and P. Heinerth, coll. PASCO COUNTY—(4) Black 
Hole, 4.0 km S of Aripeka (SW'4, NE%4, Sec. 14, T. 24S, R. 16E), 1 2, PH, 
coll.; (5) Arch Sink, 6.3 km E of U.S. Hwy. 19 at junction of Kelly Rd. and 
Aubrey Rd., (NW'4, NE4, SW", Sec. 2, T. 24E, R. 17E), 2 6 I, 1j2, 11 January 
1983, ATL, PH, coll.; (6) Nexus Sink in Beacon Woods System, 2.5 km E of 
junction of U.S. Hwy. 19 and State Rte. 52 (Bayonet Point) (NE4, SE%4, SE, 
Sec. 3, T. 25S, R. 16E), | d II, 1 2, 11 January 1983, ATL, PH, coll. Two males, 
form II, with carapace lengths of about 25 mm, are being maintained alive at the 
Smithsonian Institution, and two others at the Florida State Museum, anticipating 
their molt to form I. 

In addition, crayfish presumably belonging to this species were observed by 
A. T. Leitheuser, W. K. Fehring, P. Heinerth and other divers in at least two 
other sinks: HERNANDO COUNTY—(1) Little Springs (=Double D’s Sink, 
Twin D’s), 1.0 km SW of junction of U.S. Hwy. 19 and State Rte. 50 (Weeki- 
wachee Springs) (NE14, NW'4, SW, Sec. 2, T. 23S, R. 17E); (2) Little Salt 
Spring, 6.0 km NW of Weekiwachee Springs on State Rte. 50 (NW'4, NE, 
NEW, Sec, 297i 22S) Re): 

All of the known localities for P. (O.) leitheuseri, including those sight records 


VOLUME 96, NUMBER 2 329 


of cave divers, occur within a karst area in the Gulf Coastal Lowlands region (as 
defined by White 1970), in southwestern Hernando and northwestern Pasco coun- 
ties. The range of the species is thought to extend from the salt water of the Gulf 
of Mexico on the west to the Brooksville Ridge on the east. The northern and 
southern limits are currently undefined, but presumably they lie between the 
Withlacoochee River and Tampa Bay. The Brooksville Ridge appears to form an 
effective barrier to the eastern expansion of the species. It may also separate P. 
(O.) leitheuseri from the closely related P. (O.) lucifugus lucifugus (Hobbs, 1940) 
which occurs on the other side of the ridge near Floral City, in southeastern 
Citrus County. This “‘ridge’’ actually consists of a series of north-south trending 
ridges that reach elevations in excess of 60 meters. Their surfaces are covered 
with sand, which, in turn, is underlain with clastic sediments of the Bone Valley 
and Alachua formations (White 1970). Thick deposits of white sand, thought to 
be old stabilized dunes, occur along the western margins of the ridge complex. 
Franz and Lee (1982) proposed that when moderate to heavy accumulations of 
unconsolidated sediments cover underlying limestones, they disrupt the flow of 
organic detritus into the aquifer and produce a severely energy-limited system 
that excludes troglobitic crayfishes. 

Variations.—The tubercles, and especially the spines, ornamenting the speci- 
mens from Arch and Nexus sinks are much more prominent than in those from 
the other localities; however, their numbers and disposition do not vary conspic- 
uously. Too, the pigment in the eyes of these specimens is much denser and 
covers a larger area than that in specimens from other localities. Among the 13 
specimens for which measurements are available, there seems to be no correlation 
between size of the crayfish and the ratio of the length of the areola to its width. 
None of the males has ratios of areola length to carapace length of less than 35.3, 
but of the eight females, in only two is the ratio greater: 35.7 and 37.2, while in 
only two of the five males is the ratio greater than 37.2. As for the ratio of areola 
length to postorbital carapace length, there seems to be no correlation with car- 
apace length. 

Relationships.—Procambarus (Ortmannicus) leitheuseri has its closest affini- 
ties with P. (O.) lucifugus and its allies and, perhaps surprisingly, shares more 
in common with P. (O.) lucifugus alachua (Hobbs, 1940) than with the geograph- 
ically more proximate nominate subspecies. In addition to possessing a black 
pigment spot in the eye, the rostral margins are usually tapering anteriorly, at 
most weakly biconvex; the postorbital ridges are provided with tubercles or spines 
in addition to that capping the anterior extremity, and the areola is comparatively 
broad, all characters shared with P. (O.) lucifugus alachua rather than with the 
nominate subspecies. It differs from the former chiefly in features of the first 
pleopod of the first form male: the caudal process is obsolete; the hump, which 
is present at the cephalic base of the cephalic process in P. (O.) lucifugus alachua 
is less prominent and bent mesially; and the cephalic process is proportionately 
longer. In addition, the areola is also broader, more so than in most of the Flo- 
ridian troglobitic members of the subgenus. 

Ecological notes.—Procambarus (O.) leitheuseri has been found in flooded 
caves at water depths between 16.7 and 69.9 meters. These caves are developed 
in the upper Eocene limestones of the Ocala Group and are apparently integral 


330 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


parts of a major conduit system that carries fresh water toward the Gulf of Mex- 
ico. Vertical shafts formed in the Suwannee Limestone (Oligocene age) connect 
caves with the surface. Shafts typically open in the bottom of water-filled sinkhole 
depressions. Some of these depressions show daily fluctuations in their water 
levels that may be related to tidal activity on the Gulf. Tides may also account 
for the observed flow reversals encountered at Eagle’s Nest. According to W. K. 
Fehring, there is a salt water layer at a depth of 27.4 meters in the Beacon Woods 
system, particularly at Nexus Sink, in Pasco County. Crayfish have been ob- 
served only in the freshwater zone above the halocline. Fehring and his associates 
are studying the waters in the cave systems along the Gulf coast. 

Divers report that a rain of silt passes through the vertical shafts from the 
surface and is deposited in thick layers on cave floors. The silt either accumulates 
under the shafts or disperses into the cave, depending on the amount of water 
movement in the system. Crayfishes are usually associated with the silt. None 
has been seen in the surface ponds or in the shafts. It is believed that the organic 
fraction of the silt is generated from biotic production in the surface ponds. 

All of the known localities for P. (O.) leitheuseri lie within 11.6 km of the Gulf 
of Mexico. Black Hole and Eagle’s Nest occur 1.2 and 1.5 km, respectively, from 
the Gulf, and their surface ponds are near sea level. Both ponds receive drainage 
from adjacent wetlands. At Black Hole, the water in the pond and in the cave is 
usually nearly black in color (hence the name) due to continual drainage of acidic 
water from surrounding red bay (Persea borbonia) dominated bayheads. Die 
Polder 2 and 3 and Beacon Woods sinks occur near the 12.1 meter contour, and 
are located in sandhills near the base of the Brooksville Ridge in a longleaf pine- 
turkey oak-wire grass association. 

At Eagle’s Nest, P. (O.) leitheuseri was collected with Troglocambarus ma- 
clanei Hobbs, and the troglobitic amphipods Crangonyx grandimanus Bousfield 
and C. hobbsi Shoemaker. 


Key to Troglobitic Members of Subgenus Ortmannicus 
(Modified from Hobbs et al. and Daniel 1977:26—27 


| Eyes with: pigment)... .oc. qe). 2 lds ne ay ee eke ee 2 
— Eyes without pigment -. ..c sees eg ee ct. ees Gad 5 
2(1):) Pigment imeyes reds 5) yeh te SE. 2 3 
Hh Pigment im eyes, black. of ei fs oivaes cde at eke oe ee eee 4 


3(2) Several cervical spines present; postorbital ridge with spines caudally; 
male with hook on ischium of fourth pereiopod opposed by tubercle 
on basis; cephalic process of first pleopod of male situated lateral to 
central projection. Female with caudally directed tuberculiform pro- 
cesses on caudal margin of sternum immediately anterior to annulus 
Veil thaliSe \as2 coe Cees ees 2 aera a orcinus Hobbs and Means, 1972 

Bu Only | cervical spine present; postorbital ridge without spines cau- 
dally; male with hook on ischium of fourth pereiopod not opposed by 
tubercle on basis; cephalic process of first pleopod of male situated 
anterior to central projection. Female lacking caudally directed tu- 
berculiform processes on caudal margin of sternum immediately an- 
terior to annulus ventralis .......... erythrops Relyea and Sutton, 1975 


VOLUME 96, NUMBER 2 331 


4(2’) Areola more than 10 times as long as wide; first pleopod of male, form 
I, with caudal process and with hump at cephalic base of cephalic 
process. (The eyes of about 50% of the intergrade population in Marion 
County, Florida, lack pigment.) ...... lucifugus alachua (Hobbs, 1940) 
4' Areola less than 10 times as long as wide; first pleopod of male, form 
I, with caudal process obsolete and lacking hump at cephalic base of 
Ce MAMI MPN OCESS rete te A asp nile eiahe sae Seales aa od leitheuseri, new species 
5(1’) Male with hook on ischium of fourth pereiopod distinctly overreaching 
basioischial articulation; female with multituberculate sternum imme- 


diately anterior tosannulus* ventralis’ "2. .0.0-.24.6 4-000 s2 ae oe 6 
5s Male with hook on ischium of fourth pereiopod not overreaching ba- 

sioischial articulation; female without tubercles on sternum immedi- 

MUS RAMS I Ole FORAMMMUNNS: Us! eects sien, oom oe cunt esd eh eek @ meme ees 7 


6(5) Postorbital ridge with spines or tubercles caudally; areola less than 20 
times as long as broad. Male with cephalic process of first pleopod 
situated lateral to central projection .... horsti Hobbs and Means, 1972 

6' Postorbital ridge without caudally situated spines or tubercles; areola 
more than 20 times as long as broad. Male with cephalic process of 
first pleopod situated cephalic to central projection .................... 

TR ee te ross a int an Cahn fieustoyagh ape ots pallidus (Hobbs, 1940) 

7(5') Rostrum narrower at base than over eye; areola more than 20 times 
as long as wide; first pleopod of male with preapical curvature of at 
least 80 degrees (see note concerning eye pigment in couplet 4) ......... 

5 0 0 01d Sieh ae RRMA te mut ot ane aE lucifugus lucifugus (Hobbs, 1940) 

He Rostrum tapering from base; areola less than 20 times as long as wide; 
first pleopod of male with preapical curvature of no more than 60 
GISMIFBSS satellite aes Sea tee ere pepe ies: artic weet franzi Hobbs and Lee, 1976 


Acknowledgments 


For collecting the specimens on which this description is based and for fur- 
nishing us data concerning the habitats occupied by them, we extend our thanks 
to D. Basile, J. Bentz, G. S. Brkich, L. Collins, W. K. Fehring, S. Fehring, T. 
Gien, P. Heinerth, J. Kasserman, A. T. Leitheuser, D. P. Manor, P. C. Meng, 
J. Stone, and C. Vilece. We are also grateful to T. A. Bowman and to J. F. 
Fitzpatrick, Jr., for their criticisms of the manuscript. 


Literature Cited 


Franz, Richard, and David S. Lee. 1982. Distribution and evolution of Florida’s troglobitic cray- 
fishes.—Bulletin of the Florida State Museum, Biological Sciences 28(3):53-78, 9 figs. 
Hobbs, Horton H., Jr. 1940. Seven new crayfishes of the genus Cambarus from Florida, with notes 

on other species.—Proceedings of the United States National Museum 89(3097):387-423, figs. 
14-22. 
—. H. H. Hobbs III, and Margaret A. Daniel. 1977. A review of the troglobitic decapod 
crustaceans of the Americas.—Smithsonian Contributions to Zoology 244:v + 183 pp., 70 figs. 
,and David S. Lee. 1976. A new troglobitic crayfish (Decapoda, Cambaridae) from peninsular 
Florida.—Proceedings of the Biological Society of Washington 89(32):383-391, | fig. 
, and D. Bruce Means. 1972. Two new troglobitic crayfishes (Decapoda, Astacidae) from 
Florida.—Proceedings of the Biological Society of Washington 84(46):393-409, 2 fig. 


332 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 
White, W. A. 1970. The geomorphology of the Florida peninsula.—Florida Bureau of Geology, 
Bulletin 51, 164 pp. 


(RF) Florida State Museum, University of Florida, Gainesville, Florida 32611; 
(HHH) Department of Invertebrate Zoology, Smithsonian Institution, Washing- 


ton, D.C. 20560. 


PROC. BIOL. SOC. WASH. 
96(2), 1983, pp. 333-338 


ATYA ABELEI, A NEW ATYID SHRIMP (CRUSTACEA, 
DECAPODA, ATYIDAE) FROM THE PACIFIC 
SLOPE OF PANAMA 


Bruce E. Felgenhauer and Joel W. Martin 


Abstract.—Atya abelei is described from a single male collected from the Pa- 
cific drainage of Panama. It is easily distinguished from its sympatric congeners 
by having acute rostral prominences, no comb setae on pereopods 1 and 2, and 
three transverse white bands on the dorsum. 


The genus Atya Leach, 1815, has recently been reviewed by Hobbs and Hart 
(1982). These authors include the American and West African representatives of 
the genus which currently includes 11 species. 

Six of the 11 species have been reported from Panama in Central America 
(Abele 1975; Hobbs and Hart 1982): Atya scabra (Leach, 1815); A. innocous 
(Herbst, 1764) [a senior synonym of Atya tenella]; A. margaritacea (A. Milne 
Edwards, 1864) [a senior synonym of Atya rivalis]; A. dressleri Abele, 1975; and 
A. crassa Smith, 1871. As a result of investigations of the biology of Panamanian 
atyids (Felgenhauer and Abele, in press) an apparently new species of Atya was 
found in the Pacific drainage. The specimen had been maintained in an aquarium 
for one and one-half years and we were alerted to the possibility that it was 
unique by its color pattern, method of feeding, and size. Below we describe the 
specimen and provide notes on its feeding behavior and color. 


Atya abelei, new species 
Figs. 14 


Material.—Holotype, 6 USNM 195335, 8.1 mm cl, 49 mm tl. Type locality: 
Panama, El Valle, Cocle Province, unnamed tributary of Rio Anton, 600 m ele- 
vation, 17 Oct 1980; coll. B. E. Felgenhauer, N. H. Williams. 

Description.—Rostrum (Fig. 3A—B) short with distinct dorsal carina extending 
to sharply acute tip; latter projecting just beyond basal antennal segment. Lateral 
prominences acute. Dorsal and ventral carinae lacking teeth. Ventral carina armed 
with several rows of long plumose setae. Carapace (Fig. 3A—B) smooth, lacking 
pubescence. Antennal spine and sharply acute pterygostomial spine present. 
Distinct doublure lacking pubescence. First 4 abdominal pleura (Figs. 1, 3G) 
broadly rounded posteroventrally; ventral margins of third and fourth pleura armed 
with several long simple setae. Fifth and sixth pleura slightly angled and blunt, 
with prominent strong setae present on ventral margin of fifth. Telson (Fig. 3D) 
armed on each side of midline with row of 7 strong spines; each row beginning 
about 14 distance from anterior margin of telson and terminating in single addi- 
tional spine at each posterolateral margin. Posterior margin with 2 strong spines 
located above one another just anterior to posterior margin, and 12 long plumose 
setae. Eyes well developed and pigmented. Preanal carina (Fig. 3F, H) long, 
terminating in bifid tooth. Antennule (Fig. 3A—B) bearing single row of 5 or 6 


334 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


x : : “Ae titan tees 3 
A 
P | a ; & 
> 


IN \ 5.0mm 


Fig. 1. Atya abelei, male holotype, lateral view. 


dark spines on distal margin of basal segment; this segment with longitudinal 
rows of simple setae on dorsal surface. Second segment about twice length of 
proximal segment and bearing single medial dark spine dorsally; distal margin 
armed with 7 or 8 such spines. Dorsum of third segment with 6 or 10 dark spines 
on distal margin. Antennae extending about %4 length of body. Scaphocerite 
reaching just beyond antennular peduncle. Third maxilliped pediform, extending 
slightly beyond antennular peduncle. Fused propodus-dactylus with several rows 
of strong serrate setae. First 2 pereopods typical for genus and similar. Propodus 
and dactylus armed with long setae modified for filtering. No serrate or comb 
setae present. Third pereopod (Fig. 3C) swollen and armed with single row of 


1.0mm 1.0mm 1.0mm 


Fig. 2. Atya abelei, male holotype, dactylus and distal parts of propodus of pereopods 3 through 
5. A, pereopod 5; B, pereopod 4; C, pereopod 3. 


VOLUME 96, NUMBER 2 335 


G 


5.0mm H 1.0mm 


Fig. 3. Atya abelei, male holotype. A, anterior region, lateral view; B, anterior region, dorsal 
view; C, pereopod 3; D, telson, dorsal view; E, appendices masculina and interna; F, preanal carina, 
lateral view; G, Second through fifth abdominal pleura; H, Sternum of sixth abdominal segment and 
preanal carina. 


336 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 4. Color patterns of Panamanian atyids. White arrow indicates Atya abelei. Black on white 
arrow points to A. innocous. Shrimp feeding in center is A. margaritacea. The larger shrimp feeding 
at bottom left is another individual of A. innocous. 


densely plumose setae originating on lateral surface of merus and continuing to 
tip of propodus. Merus with 2 strong movable spines on ventromesial side. Short 
sclerotized spines present on dorsal surface of merus, carpus, and propodus. 
Merus approximately 6 times as long as wide, about 3 times length of carpus, 
slightly more than twice length of propodus, and about 7 times length of dactylus. 
Flexor surface of dactylus (Fig. 2C) bearing single row of 5 blunt spines; distal 
part narrow and acute. Fourth pereopod similar to third but less swollen. Merus 
with 2 movable spines on ventrolateral margin and 3 sharp spines on correspond- 
ing part of carpus. Dactylus (Fig. 2B) bearing row of 5 strong spines on flexor 
surface; distal part narrow and acute. 

Fifth pereopod similar to fourth with single short sharp spine on ventrolateral 
margin of ischium-basis. Merus armed with 3 sharp movable spines on ventrome- 
sial margin. Carpus with | large sharp movable spine on corresponding surface. 
Dactylus (Fig. 2A) armed differently from that in other pereopods, bearing single 
row of approximately 25 serrate denticles on lateral flexor margin; tip sharp and 
acute. Endopod of first male pleopod broadly ovate, subequal in length to exopod, 
with scattered short curved spines on anterior surface. Appendix masculina (Fig. 
3E) slightly more than 3 times as long as wide. Margins with long curved spines, 
mesial surface bearing scattered long spines. Proximal region armed with several 
long plumose setae. 

Color.—The color pattern described below is from the single live male kept in 
an aquarium at Florida State University. The ground color of the ventral half of 
the body is light grey and the upper half rust brown. The intensity of the brown 


VOLUME 96, NUMBER 2 337 


dorsum increases anteriorly. Light yellow specks are present on all parts of the 
body. The light brown carapace is marked with a prominent transverse band 
distinctly bordered posteriorly by black. On the upper posterior surface of the 
carapace there is a black band that decreases in width ventrally. 

The first abdominal segment has a white transverse band covering much of the 
segment. The third and fourth abdominal segments are light brown. The fifth 
segment has a small white band on its dorsal surface, and the sixth is entirely 
white. The proximal region of the uropods is also white and in the distal region 
blending to yellowish brown. The antennular peduncle and antenna are concol- 
orous light brown. The pereopods and pleopods are all translucent brown. 

Etymology.—This new species is named in honor of Lawrence G. Abele for 
his many contributions to crustacean biology. 

Habitat.—Atya abelei inhabits rock rubble in water flowing at a velocity of 
1.5-2.0 m/sec. Atya abelei and A. margaritacea occur in the same microhabitat 
with A. innocous, primarily found clinging to vascular plants at the stream edge 
(Felgenhauer and Abele, in review). 

Feeding Behavior.—Atya abelei feeds primarily by passive filtration. Careful 
observations of this shrimp in aquaria for over a year revealed little evidence for 
scraping the substrate for food as is common for many atyids. The absence of 
scraping setae on the chelate pereopods would inhibit the shrimp’s ability to 
remove material effectively from the substrate and probably accounts for the 
minimum amount of scraping in this species. 

Discussion.—Atya abelei appears most closely related to Atya innocous which 
occurs in both the Atlantic and Pacific drainages of Panama and the West Indies 
(Abele and Blum 1977; Chace and Hobbs 1969). The species can easily be distin- 
guished in that A. abelei does not have scraping denticles on the terminal brushes 
of the first and second pereopods. In addition, the angle of the lateral prominences 
of the rostrum is acute in A. abelei and the preanal carina terminates in a bifid 
tooth; these conditions are never present in specimens of A. innocous (Hobbs 
and Hart 1982). Other features that serve to separate the two species include the 
following: the ventral margin of the second through the fifth abdominal pleura in 
some populations of A. innocous is armed with sclerotized denticles on the ventral 
margins (Hobbs and Hart 1982), whereas A. abelei lacks these denticles and 
instead exhibits long simple setae in this region; the merus of the third pereopod 
of A. innocous is unarmed while the merus of A. abelei has two strong movable 
spines on the inner margin. In addition to the above characters, the species can 
be distinguished in the field by its color (Fig. 4). Atya innocous exhibits a lon- 
gitudinal stripe extending from the base of the rostrum to the anterior margin of 
the telson. Atya abelei has large white transverse bands bordered anteriorly or 
posteriorly in black on the dorsum of the carapace, first abdominal segment, and 
fifth abdominal segment. 


Acknowledgments 


We wish to thank Dr. N. H. Williams for assistance in the field. Special thanks 
must be given Dr. Robert Dressler and the Smithsonian Tropical Research Insti- 
tute for their kind hospitality and use of facilities in Panama. We are extremely 
grateful to Drs. Horton H. Hobbs, Jr., and C. W. Hart, Jr., for allowing us to see 


338 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


a manuscript xerox copy of their monograph on the genus Atya. Lastly, we must 
thank Dr. Horton H. Hobbs, Jr., for critically commenting on an early draft of 
the manuscript. Funds were provided by the National Science Foundation (Doc- 
toral Improvement Grant) NSF No. DEB-801835. 


Literature Cited 


Abele, L. G. 1975. A new species of freshwater shrimp (genus Atya) from the Pacific drainages of 

Panama.—Proceedings of the Biological Society of Washington 88:51—58. 

,and N. Blum. 1977. Ecological aspects of the freshwater decapod crustaceans of the Perlas 

Archipelago, Panama.—Biotropica 9:239-252. 

Chace, F. A., Jr., and H. H. Hobbs, Jr. 1969. The freshwater and terrestrial decapod crustaceans 
of the West Indies with special reference to Dominica.—Bulletin of the United States National 
Museum 292: 1-258. 

Felgenhauer, B. E., and L. G. Abele. (In press). Aspects of mating in the tropical freshwater shrimp 
Atya innocous (Herbst).—Biotropica. 

, and Ultrastructure and ecological morphology of the feeding and associated ap- 

pendages of the tropical tropical freshwater shrimp Atya innocous (Herbst). (In review). 

Hobbs, H. H., Jr., and C. W. Hart, Jr. 1982. The shrimp genus Atya (Decapoda: Atyidae). Smith- 
sonian Contributions to Zoology 364: iii + 143 pp. 


Department of Biological Science, Florida State University, Tallahassee, Flor- 
ida 32306. 


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INFORMATION FOR CONTRIBUTORS 


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CONTENTS 


Three new species of Colobomatus (Copepoda: Philichthyidae) parasitic in the mandibular canals 
of haemulid fishes Roger F. Cressey and Marilyn Schotte 
Notes on the biology of some seagrass-dwelling crustaceans (Stomatopoda and Decapoda) from 
Caribbean Panama Loren D. Coen and Kenneth L. Heck, Jr. 
Studies of neotropical caddisflies, XXXIV: the genus Plectromacronema (Trichoptera: 
Hydropsychidae) Oliver S. Flint, Jr. 
A revision of the Bogueidae Hartman and Fauchald, 1971, and its reduction to Bogueinae, a 
subfamily of Maldanidae (Polychaeta) Paul S. Wolf 
A new species of Cancellaria (Mollusca: Cancellariidae) from the northern Gulf of Mexico 
Richard E. Petit 
A new species of polymorphic fish, Cichlasoma minckleyi, from Cuatro Ciénegas, Mexico (Te- 
leostei: Cichlidae) Irv Kornfield and Jeffrey N. Taylor 
Clarification of the names Rana mystacea Spix, 1824, Leptodactylus amazonicus Heyer, 1978 
and a description of a new species, Leptodactylus spixi (Amphibia: Leptodactylidae) 
W. Ronald Heyer 
A new bathyal species of Coralliodrilus (Oligochaeta: Tubificidae) from the southeast Atlantic 
Christer Erséus 


New species of Fabriciola and Fabricia (Polychaeta: Sabellidae) from Belize Kirk Fitzhugh » 


Bermudalana aruboides, a new genus and species of troglobitic Isopoda (Cirolanidae) from marine 
caves on Bermuda Thomas E. Bowman and Thomas M. Iliffe 
Varichaetadrilus, a new name for Varichaeta Brinkhurst, 1981, non Speiser, 1903, (Diptera) 
with a description of a new species V. fulleri 
Ralph O. Brinkhurst and R. Deedee Kathman 
Bioluminescence in the marine ostracod Cypridina americana (Miller, 1890) off Manzanillo, 
Mexico (Myodocopa: Cypridininae) David Lapota 
Eschmeyer nexus, a new genus and species of scorpaenid fish from Fiji 
Stuart G. Poss and Victor G. Springer 
Fennerosquilla, a new genus of stomatopod crustacean from the northwestern Atlantic 


Raymond B. Manning and David K. Camp | 


Procambarus (Ortmannicus) leitheuseri, new species, another troglobitic crayfish (Decapoda: 
Cambaridae) from peninsular Florida Richard Franz and Horton H. Hobbs, Jr. 
Atya abelei, a new atyid shrimp (Crustacea, Decapoda, Atyidae) from the Pacific slope of 
Panama Bruce E. Felgenhauer and Joel W. Martin 


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Proceedings 

of the 

_ BIOLOGICAL SOCIETY 

: ‘ 
WASHINGTON 


31 October 1983 Number 3 


THE BIOLOGICAL SOCIETY OF WASHINGTON 


1983-1984 
Officers 
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Custodian of Publications: Michael J. Sweeney 


PROCEEDINGS 
Editor: Brian Kensley 


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PROC. BIOL. SOC. WASH. 
96(3), 1983, pp. 339-348 


NECTOCARMEN ANTONIOI, A NEW PRAYINAE, 
CALYCOPHORAE, SIPHONOPHORA 
FROM CALIFORNIA 


Angeles Alvarino 


Abstract.—Nectocarmen antonioi n. gen., n. sp. was found 13 April 1981 at 
33°36.7'N, 118°18.4'W, off California, at a depth of less than 70 m. The siphono- 
phore was complete, with nectophores, gastrozooids, tentacles, palpons, tentilla, 
bracts, and gonophores. It differs from other genera (Rosacea, Nectodroma or 
Praya, Prayoides, Lilyopsis, Desmophyes, Stephanophyes) in the subfamily 
Prayinae in the characteristics of its nectophores, bracts, and gonophores, but 
constitutes a morphological link among them. 


Nectocarmen, new genus 


Diagnosis.—A member of the subfamily Prayinae, which includes the genera 
Rosacea (2 species), Nectodroma or Praya (2 species), Lilyopsis (2 species), and 
Prayoides, Desmophyes, and Stephanophyes (1 species each). The principal char- 
acteristics of these genera and of Nectocarmen are given in Table 1. 

Type-species.—Nectocarmen antonioi, new species. 

Etymology.—Necto (“‘swimming,’’ Greek) + Carmen, my mother’s name (Latin 
for poetry, song, and garden). 


Nectocarmen antonioi, new species 
Figs. 1—5 


Material.—Off California, San Pedro Channel, 33°36.7'’N, 118°18.4’W, R/V Da- 
vid Starr Jordan sta 88.3 33, cruise 8104, 13 April 1981, vertical tow from about 
70 m to surface: Holotype, complete colony, USNM 60895; paratype, eudoxids, 
USNM 60896. 

Etymology.—Named for my father, Antonio. 

Description.—The subfamily Prayinae of the suborder Calycophorae are large 
prayids with pulsating nectophores and a heavy stem. The bracts are composed 
of a large amount of mesoglea, which aids the flotation of the whole animal. 
Prayids lack the apical gas-filled float of the Physonectidae, and are propelled by 
the nectophores at the upper part of the stem, which also supports the large 
bracts, gastrozooids, palpons, tentacles, tentilla, and gonophores. 

The holotype is based ona polygastric specimen with a pair of small nectophores 
at the top of the stem, and well-developed eudoxids hanging from the stem, along 
with bracts, gastrozooids, tentacles, palpons, tentilla, gonophores, and free cor- 
midia. The details of the siphonophore observed have been carefully reproduced, 
and the whole animal represented in detail in Fig. 1. Drawings give a more 
reliable representation of specimens than photographs. 

Polygastric phase: In the polygastric phase, Nectocarmen antonioi exhibits 2 
transparent kidney-shaped cylindrical nectophores, one slightly larger than the 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


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342 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 1. Nectocarmen antonioi, complete specimen, about 200 mm long. 


VOLUME 96, NUMBER 3 343 


Fig. 2. Pair of nectophores of Nectocarmen antonioi (3-4 mm long): A, Small nectophore; B, 
Large nectophore; C, Dorsal view of either nectophore. Hydroecium, nectosac, radial and circular 
canals, and somatocyst can easily be observed. 


other, with a large nectosac reaching nearly to the top of the nectophore. The 
nectophores are 3-4 mm long and about 2.5 mm wide. The hydroecium is only 
1’ the width of the nectophore, and does not run its entire length; it forms a deep 
groove, with rounded lobes (superior and inferior) closing the entrance, and this 
whole structure is covered by the flaps of the nectophore walls. 

The pair of nectophores are bound together by the hydroecium, embracing 
each other by this structure (Fig. 2). 

The nectosac is large, similar to that of Stephanophyes superba Chun, 1885 
(1891) but with differences in the hydroecium, somatocyst, and radial canals. The 
pallial canal extends slightly up the nectosac, forming the radial canals. The 
anterior and posterior radial canals are straight, and the lateral canals bend up- 
ward to a position close to the top of the nectosac, then turn straight down to 
the ring or circular canal. The velar end of the nectosac, the ostium, is large, 
extending upward at the dorsal side. No ring of minute pigment flecks or tubercles 
is observed on the ostium. 

The somatcyst is pear-shaped, and lies vertically in the mesoglea, extending 
slightly above the top of the nectosac. 

Gastrozooids retain the larval tentacle, modified to form a series of tentilla. 
The base of the gastrozooids is enlarged into the nematocyst-producing area, the 
basigaster. 

Palpons of reduced gastrozooids, with simple tentacles or palpacles, serve as 
feelers or testers. 

The tentilla, with the subterminal battery of nematocysts, are spirally twisted 
filaments, with cnidobands (sabre-shaped nematocysts) forming the chief part of 
the stinging organs. 

Gastrozooids, tentacles, palpons and tentilla lie under the bracts, and are shown 
in Fig. 1, and in detail in Fig. 3. 


344 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON . 


Fig. 3. Gastrozooids of Nectocarmen antonioi (2—3 mm long): A, Tentacles; B, Tentilla. 


Eudoxid phase: The eudoxid phase has no asexual nectophore. 

The bracts, which reach a maximum length of 10-12 mm, are oval, kidney- 
shaped, laterally flattened masses of mesoglea, the inferior part of which is con- 
cave and the upper part convex. They are divided into 2 lobes, with a third lobe 
on the concave distal side forming part of the hydroecial cavity of the bract. 
Unlike Desmophyes, no central organ (pear-shaped vesicle) is present at the 
junction of the main bracteal canals. 

The bracts contain 4 large canals. The dorsal bracteal canal, which is slightly 
curved, ends in an ampullar dilatation, as does the nearly straight ventral bracteal 
canal; left and right hydroecial canals also end in ampullar dilatations. The right 


VOLUME 96, NUMBER 3 345 


A 


Fig. 4. Bracts of Nectocarmen antonioi (10-12 mm longest axis): A, Lateral right view of two 
bracts; B, Lateral left view of two bracts. 


hydroecial canal is longer than the left. The former curves upwards and then 
extends down, and the latter bends down and then up. Usually, the left canal is 
straighter than the right. Neither the bracteal nor the hydroecial canals branch 
into small branchlets or spurs (Fig. 4). 

The bracts, with gastrozooids, tentacles, palpons, tentilla and gonophores, may 
detach from the stem and swim freely in the ocean. 

The sexual gonophores are large and hemispherical in shape, laterally flattened, 
retaining the medusoid characteristics, but without marginal tentacles. Female 
and male gonads occupy the mouthless manubrium. The gonophores pulsate in- 
termittently, and are capable of propelling the cormidia in the water. 

The gonophores are budded at the stem, below the base of the gastrozooids. 
They are, as explained, large bells, laterally compressed, with 4 straight radial 
canals ending at the ring canal. The bells hang from the bract by the pedicular 
canal, and the four radial canals run from the pedicel to the circular canal. Near 
the top of the umbrella, the pedicel has trifid branches embedded in the mesoglea, 
as 1s seen in Nectodroma (Praya) reticulata (Bigelow, 1931). The large, pear- 
shaped vesicles inside are the female gonads, containing numerous small ova. 
Male gonads are small, pointed, sausage-shaped vesicles (Fig. 5). 

Male and female eudoxids may be found on the same stem. 

It is probable that each cormidium includes either male or female gonophores, 
providing a sexual separation when male cormidia and female cormidia swim 


346 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 5. Gonophores of Nectocarmen antonioi (3-4 mm high): A, Female gonophores; B, Male 
gonophore; C, Upper view of gonophore to show the trifid branching pedicel. 


freely in the ocean. Some of the cormidia were detached, but obviously belonged 
to the same whole animal. 

The nectophores, bracts, and gonophores are transparent; gastrozooids, pal- 
pons, and tentacles range in color from bright reddish to faded pink or rose. 

The nectophores function in propulsion; the bracts are composed of enlarged 
masses of mesoglea and act as protection for gastrozooids and gonozoids or 
gonophores, as well as contributing to the buoyancy of the animal. The stem 
contracts and expands, and the gonophores also perform active pulsations, con- 
tributing to the propulsion of the siphonophores; by spreading the stem into loops 
they also amplify the potential field of the tentacles and the fishing predatory 
capacity of the siphonophore. 

Discussion.—The principal differential characteristics of the genera in the 
subfamily Prayinae are presented in Table 1, together with those for Nectocar- 
men n. gen. 

The general shape of the nectophores is similar to that of the other genera in 
the subfamily, especially Rosacea Bigelow, 1911, and Desmophyes Haeckel, 1888, 
also described by Kawamura (1915). However, the hydroecium is smaller than that 
of Rosacea, or Desmophyes, and similar to that of Stephanophyes. The nectosac 
is large, as in Stephanophyes and Lilyopsis (Alvarino 1981; Chun 1885, 1891), 
but the radial canals do not display the meandering pattern observed in Stepha- 
nophyes and Lilyopsis. The somatocyst of Nectocarmen is different from that of 
other Prayinae genera, including Prayoides intermedia Leloup, 1934. 


VOLUME 96, NUMBER 3 347 


The bracts resemble in general outline those of Nectodroma (Praya), but differ 
in the detail of the shape and pattern of the canals, as can be observed by com- 
paring present illustrations with those by Alvarino (1981), Bigelow (1911), and 
Totton and Bargmann (1965). Bracts in Nectocarmen antonioi do not have the 
vesicular organ of Desmophyes. 

The eudoxids of the new siphonophore do not have the large asexual eudoxid 
nectophore that is found in Lilyopsis, Desmophyes, and Stephanophyes. The 
gonophores are similar to those of Nectodroma (Praya); however, sexual necto- 
phores are pear-shaped in Nectodroma, and hemispherical in Nectocarmen. Both 
Nectodroma (Praya) and Nectocarmen present the trifid branched pedicular ca- 
nal, but the gonads of the former include 4—6 large ova (Bigelow 1931), while in 
the latter the ova are small and numerous. In Nectocarmen the four radial canals 
of the gonophore converge at the pedicel, and run straight to the circular canal, 
characteristics different from the other genera in the subfamily, though similar to 
Nectodroma (Praya). No tubercles or pigmented spots appear on the margin of 
the gonophoral umbrella, whereas in the other species of the subfamily those 
structures are present. 

Species of the following Prayinae have been observed in California waters 
(Alvarino 1967, 1971): Nectodroma dubia (Quoy and Gaimard, 1833), N. retic- 
ulata (Bigelow, 1911), Rosacea plicata Bigelow, 1911. Bigelow and Leslie (1930) 
reported N. reticulata from Monterey, but Totton and Bargmann (1965) indicate 
it was probably N. dubia. 

There is a peculiar phenomenon common to Siphonophorae and Medusae, in 
that most of the species present an erratic distribution (Alvarino 1977, 1981; Sears 
1953). Sears (1953:11) says: ‘“‘There appears to be a tendency among coelenterates 
for a seemingly good species to appear in a particular locality, often in consid- 
erable numbers, and after a time to disappear, never to be seen again.’ This 
must be related to the peculiarity in patchiness and irregular aggregational dis- 
tribution characteristics of siphonophores and other coelenterate populations, and 
the ineffectiveness of the plankton sampling gear in capturing these organisms 
(Biggs, Bidigare, and Smith 1981; Hamner eft al. 1975). Because of their swimming 
behavior and speed, and their life cycle characteristics, siphonophores are ca- 
pable of avoiding capture by plankton nets (Biggs 1977). 


Acknowledgments 


I would like to express my appreciation to Drs. Izadore Barrett, Frank Ferrari, 
John R. Hunter, and Reuben Lasker for reading the manuscript, and to Dr. 
Thomas E. Bowman and Ms. Martha Brown for their careful editing of the paper. 
My thanks are also due to Esperanza Manaligod for typing the table included in 
this work. 


Literature Cited 


Alvarifio, A. 1967. Bathymetric distribution of Chaetognatha, Siphonophorae, Medusae and Ctenoph- 
ora off San Diego, California—Pacific Science 21:473—-485. 

—. 1971. Siphonophores of the Pacific, with a review of the world distribution.—Bulletin 
Scripps Institution of Oceanography, University of California San Diego, La Jolla, California 
16: 1-432. 

—. 1977. Indicadores planctonicos: Distribucion batimetrica de algunas Medusas (Planktonic 


348 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


indicators: Bathymetric distribution of some Medusae).—Memorias II Simposio Latinoame- 

ricano sobre Oceanografia Biologica, Venezuela 1975 1:161—190. 

1981. Siphonophorae. Atlas del Zooplancton del Atlantico Sud-occidental—Publication In- 
stituto Nacional de Investigacion y Desarrollo Pesquero [INIPED], Ministerio de Comercio e 
Intereses Maritimos, Subsecretaria de Intereses Maritimos, Argentina, pp. 383-441. 

Bigelow, H. B. 1911. Siphonophorae. ALBATROSS expedition to the eastern tropical Pacific.— 
Memoirs of the Museum of Comparative Zoology at Harvard College 38(2):173—401. 

—. 1931. Siphonophorae from the ARCTURUS oceanographic expedition.—Zoological Sci- 

entific Contribution, New York Zoological Society 8(11):525—592. 

, and M. Leslie. 1930. Reconnaissance of the waters and plankton of Monterey Bay, 1928.— 

Bulletin of the Museum of Comparative Zoology at Harvard College 70:492-581. 

Biggs, D. C. 1977. Field studies of fishing, feeding and digestion in Siphonophores.—Marine Be- 

haviour and Physiology 4(4):261—274. 

, R. R. Bidigare, and D. E. Smith. 1981. Population density of gelatinous macrozooplankton: 

In situ estimation in oceanic surface waters.—Biological Oceanography 1(2):157—173. 

Chun, C. 1885. Ueber die cyclische Entwickelung der Siphonophores.—Sitzungsberichte der kon- 
glich Preussischen Akademie der Wissenschaften zu Berlin 1885:511—529. 

. 1891. Die Canarischen Siphonophores in monographischen Darstellungen I. Stephanophyes 
superba und die familie der Stephanophyiden.—Abhandlungen senckenbergische naturfor- 
schende Gesellschaft 16:553-627. 

Haeckel, E. 1888. Report on the Siphonophorae.—Report on the Scientific Research Expedition, 
H.M.S. CHALLENGER, Zoology 21:1—380. 

Hamner, W. M., L. P. Madin, A. L. Alldredge, R. W. Gilmer, and P.O. Hamner. 1975. Underwater 
observations of gelatinous zooplankton: Sampling problems, feeding biology, and behavior.— 
Limnology and Oceanography 20(6):907-917. 

Kawamura, T. 1915. Calycophorae, II.—Zoological Magazine, Tokyo 27:317-324. 

Leloup, E. 1934. Siphonophores de |’Ocean Atlantique tropical et austral—Bulletin Muséum Royal 
Histoire Naturelle de Belgique 10(6): 1-87. 

Sears, M. 1953. Notes on Siphonophores. 2. A revision of the Abylinae.—Bulletin of the Museum 
of Comparative Zoology at Harvard College 109(1): 1-119. 

Totton, A. K., and H. E. Bargmann. 1965. A synopsis of the Siphonophora.—Bnitish Museum, 
Natural History, pp. 1-230. 


National Marine Fisheries Service, NOAA, Southwest Fisheries Center, P.O. 
Box 271, La Jolla, California 92038. 


PROC. BIOL. SOC. WASH. 
96(3), 1983, pp. 349-354 


A NEW STROMBINA SPECIES 
(GASTROPODA: PROSOBRANCHIA) FROM THE 
TROPICAL WESTERN ATLANTIC 


Richard S. Houbrick 


Abstract.—Strombina (Cotonopsis) argentea is the fourth living Strombina 
species and the first Cotonopsis taxon described from the tropical western At- 
lantic. It has the most northern distribution of any western Atlantic Strombina 
species and occurs in deep water. This species is the largest of the four Atlantic 
taxa, and morphologically resembles the eastern Pacific Strombina (Cotonopsis) 
deroyae Emerson and D’ Attilio, 1969. There is considerable intraspecific varia- 
tion in axial sculpture. The radula is typically columbellid in form. 


While processing material dredged by the National Marine Fisheries ship R/V 
Oregon in deep water near Silver Bank off the north coast of the Dominican 
Republic, ten specimens of a new, remarkably large Strombina species were 
recognized. Two were live-collected and contained dried animals, allowing ex- 
amination of the operculum and radula. 

The genus Strombina was widespread in the Caribbean during the Miocene, 
but the diversity of this taxon decreased in the Pliocene. Although there are many 
species living in the eastern Pacific today, only three Recent species were pre- 
viously known from the western Atlantic, all from the southern Caribbean, and 
referred to the subgenus Strombina s.s. This paper describes a fourth Strombina 
species that is allocated to the subgenus Cotonopsis Olsson, a group previously 
known only from the Neogene of Central America and the Recent eastern Pacific 
fauna. The presence of a large, distinctive Strombina living in deep water to the 
north of the Greater Antilles and Caribbean Sea adds significant dimension to the 
composition and distribution of this lineage in the western Atlantic. 


Description 


Family Columbellidae 
Strombina Morch, 1852 
Subgenus Cotonopsis Olsson, 1942 
Strombina (Cotonopsis) argentea, new species 
Figs.. 1-2 


Shell (Table 1).—Shell large and slender, fusiform, with elongate tapering spire 
comprising 10 weakly inflated whorls and ranging in length from 30-43 mm. Whorls 
sculptured with 19-28 weak slightly curved axial ribs extending length of whorl. 
Some shells nearly smooth but early whorls of teloconch sculptured with distinct 
axial ribs. Protoconch large, bulbous, smooth, about 1!2 whorls. Body whorl 
large, nearly smooth, round in cross section, slightly angulate at periphery, about 
one-half shell length. Axial ribs present on anterior of body whorl but fading out 
near lip. Thick varix just above edge of outer lip. Siphonal constriction of body 


350 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


F 


Fig. 1. Strombina (Cotonopsis) argentea (all specimens whitened with ammonium chloride to 
enhance sculptural details—Fig. G is natural; A, B, C, Apertural, side and dorsal views of holotype, 
USNM 810345, 39 mm; D, Operculum of holotype, 6 mm long; E, Detail of protoconch and sculpture 
of early whorls, USNM 821851, 22.5x; F, Paratype, showing protoconch, USNM 821851, 31 mm; 
G, Dorsal view, same as F, but natural to show glossy shell and light tan color pattern; H, Weakly 
sculptured specimen, paratype USNM 821851, 43 mm; I, Specimen with strong axial sculpture, para- 
type, USNM 821851, 41 mm. 


VOLUME 96, NUMBER 3 351 


Fig. 2. Scanning electron micrographs of radula of Strombina (Cotonopsis) urgentea. Lateral teeth 
spread back to show weak rachidian teeth. Note large interspaces on radular membrane between 
rachidian and lateral teeth. 


whorl sculptured with fine spiral striae. Suture straight, slightly impressed. Ap- 
erture little less than one-half shell length, narrow, lunate, sinuous. Anterior 
siphonal canal long, curved to left, slightly reflected dorsally. Anal canal short, 
deeply incised. Parietal area somewhat thickened. Columella concave, sinuous 
and twisted at canal constriction. Slight columellar wash present. Outer lip smooth 
with thickened sinus at anal canal and slight posterior extension onto body whorl. 
Inner lip with tiny denticles and | to 2 larger teeth adjacent to anal canal. Denticles 
not extending into aperture. Shell color white with 2 broad bands of faded, yel- 
lowish-tan zigzag markings. 

Operculum (Fig. 1, D).—Operculum thin, corneous, lenticular with terminal 
nucleus at edge. 

Radula (Fig. 2, Table 2).—Radular ribbon rachiglossate (1+1+1), typically 
columbellid in form, narrow, relatively long, little over one-fourth shell length, 
comprising about 245 rows of teeth. Rachidian tooth a weak, narrow, nearly 
rectangular plate with rounded dorsal edges, pointed where joined to basal mem- 
brane. Wide interspace between rachidian tooth and lateral teeth. Lateral tooth 
comprising a shaft with peg-like base, 2 hooked cusps at tip, single blunt hook 
on mid shaft of tooth. 

Animal.—Preserved, dried animals, when rehydrated, appeared to be unpig- 
mented and had a large foot, and a smooth mantle edge with a muscular, well- 
developed anterior siphon. Head small, with short tentacles each with a small 
basal eye. Proboscis pleurembolic, long, containing large buccal mass and long 
radular ribbon. The poor state of preservation did not allow detailed anatomical 
studies. 


Table 1.—Shell measurements of Strombina argentea (all measurements in mm). 

Statistic (n = 10) ve SD Var. Range 
Length 38.83 3.57) 11.45 30-43 
Width 12.97 1.26 1.42 10.3-14.6 
Length of aperture 18.05 1.95 3.43 14—20.6 
Width of aperture 5.21 0.58 0.31 4.1-6.2 


No. of whorls 10.1 0.32 0.09 10-11 


B52 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Table 2.—Radular measurements (all measurements in mm). 


Shell length 
Character (n = 2) Radula length Rows of teeth Shell length Radula length 
Mean 10.2 244.8 39.5 4.07 


Type-locality.—384—430 m, 20°48'N, 70°46'W, near Silver Bank, 60 miles NE 
of Luperon, Dominican Republic; NMFS, R/V Oregon, Station 5432. 

Holotype (Fig. 1, A-B).—USNM 810345; length 39 mm, width 13.1 mm. 9 
paratypes, USNM 821851 (for measurements see Table 1). 

Etymology .—From the Latin adjective argenteus, “‘silver,’’ in reference to the 
Silver Bank area, where the specimens were taken. 


Remarks 


Specimens of this species were dredged in depths of 210—235 fms (384-430 m) 
in the Silver Bank Passage between Mouchoir Bank and Silver Bank. The shell 
form and axial sculpture are adapted to a sandy bottom habitat and all dead shells 
had drill holes indicative of predation by naticid snails. Several shells showed 
repair marks due to crab attacks. Two specimens were collected alive. Within 
the rectum of one of these were the remains of a sipunculan worm of the genus 
Aspidosiphon. 

The degree of axial sculpture shows considerable intraspecific variation. Some 
Shells are nearly devoid of axial ribs while in others they may be pronounced. 
Axial sculpture is strongest on the early, post-nuclear whorls and on the penul- 
timate and body whorls, and is most pronounced on the anterior portion of each 
whorl, adjacent to the suture. When axial ribbing is strong, the suture appears 
wavy. 

Among the Recent Panamic fauna, Strombina deroyae Emerson and d Attilio, 
1969, from the Galapagos, assigned by Keen (1971) to Cotonopsis, is morpho- 
logically close to the western Atlantic Strombina argentea although the color 
pattern of the latter is weaker. Strombina argentea also resembles Strombina 
fusinoidea Dall, 1916, in general form, but that taxon is a Strombina s.s., and 
differs accordingly. 

Strombina argentea has a shell remarkably convergent with that of Cyomesus 
chaunax (Bayer, 1971), a totally unrelated turbinellid species. The latter differs 
chiefly in having strong columellar plications. 

Strombina is not a common taxon in the Recent Caribbean fauna. Olsson and 
Harbison (1953:230) pointed out that Strombina is characteristically Panamic in 
the Recent fauna, but in the Miocene and Pliocene it was well represented in an 
extended Caribbean province by a number of species. It also occurred in north 
Florida (Petuch 1982:304). Petuch (1980:86) noted that there are only three living 
species in the western Atlantic, while Keen (1971) recorded 25 species living in 
the eastern Pacific. Weisbord (1962:327—328) listed about 25 fossil species from 
the Caribbean and as many as 40 species are named from Neogene deposits in 
tropical America (Woodring 1964:252). J. and W. Gibson-Smith (1974), in an 
extensive overview of this group in the Caribbean, estimated about 60 fossil taxa 
occurred in that province during Miocene and Pliocene times. 


VOLUME 96, NUMBER 3 353 


Three subgenera of Strombina have been proposed: Strombina s.s., Cotonopsis 
Olsson, 1942, and Sincola Olsson and Harbison, 1953. Although most western 
Atlantic living and fossil species are allocated to Strombina s.s., J. and W. Gib- 
son-Smith (1974:51) pointed out several Caribbean fossil taxa which do not fit the 
limits of these subgenera and suggested that the entire group was in need of 
revision. 

The three living Caribbean species, Strombina pumilio (Reeve, 1859), S. ca- 
boblanquensis Weisbord, 1962, and S. francisiae J. and W. Gibson-Smith, 1974, 
are all assigned to Strombina s.s. None of these taxa bears any resemblance to 
the much larger Strombina argentea, which falls within the limits of the subgenus 
Cotonopsis. Cotonopsis differs from Strombina s.s. in lacking a thickening or 
hump on the back of the body whorl, by the circular section of the body whorl, 
a less thickened lip that is not strongly denticulate within, and in the more 
strongly recurved anterior canal. Cotonopsis was originally based on two fossil 
species in the Tertiary beds of the Burica Peninsula, Panama. Keen (1971) has 
assigned four living eastern Pacific species to Cotonopsis. The subgenus Coto- 
nopsis was previously unrecorded in the Recent or fossil fauna from the western 
Atlantic. A review of the Caribbean fossils depicted in the literature revealed 
nothing similar to Strombina argentea; moreover, none of the fossils from the 
Dominican Republic shown by Maury (1917, pl. 15) and Pilsbry (1921) or those 
of Jamaica (Woodring 1928) are similar to this species. Exclusive of S. argentea, 
all Recent and most fossil species from the western Atlantic belong to Strombina 
s.s. The type-species of Cotonopsis, Strombina panacostaricensis Olsson, 1942, 
has axial ribs on each whorl and the spiral striae on the body whorl constriction 
seen in S. argentea, but lacks the long twisted anterior canal of the latter. 


Acknowledgments 


I wish to thank Dr. M. J. Harasewych for his help with photography and for 
critically reading the manuscript. My thanks to Ms. J. Piraino for taking the SEM 
micrographs and to Mrs. June Jones for typing the drafts of this paper. Dr. Mary 
Rice identified the sipunculan remains. This is contribution No. 115 of the Smith- 
sonian Marine Station, Ft. Pierce, Florida. 


Literature Cited 


Bayer, F. M. 1971. New and unusual mollusks collected by R/V John Elliott Pillsbury and R/V 
Gerda in the tropical western Atlantic.—Bulletin of Marine Science 21(1):111—236. 

Emerson, W. K., and A. D’ Attilio. 1969. A new species of Strombina from the Galapagos Islands.— 
Veliger 11(3):195—197, pl. 39. 

Gibson-Smith, J., and W. Gibson-Smith. 1974. The genus Strombina (Mollusca: Gastropoda) in 
Venezuela, with descriptions of a new Recent and some fossil species.—Boletin Informatino. 
Associacion Venezolana-de Geologia, Mineria y Petroleo 17(4,5,6):49-70, pls. 1-4. 

Keen, M. A. 1971. Sea shells of tropical west America, 2nd Ed. Stanford. 1064 pp., 22 pls. 

Maury, C. J. 1917. Santo Domingo type sections and fossils —Bulletins of American Paleontology 
5(29): 1-249, 39 pls. 

Olsson, A. A., and A. Harbison. 1953. Pliocene mollusca of southern Florida.—The Academy of 
Natural Sciences of Philadelphia, Monograph No. 8, 456 pp., 65 pls. 

Petuch, E. J. 1980. A reanalysis of Neogene Caribbean provinciality with reference to the discovery 
of a relict Caenogastropod fauna off northern South America. Unpublished Ph.D. Dissertation, 
University of Miami, 163 pp. 


354 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


—.. 1982. Geographical heterochrony: contemporaneous coexistence of Neogene and Recent 
molluscan faunas in the Americas.—Palaeogeography, Palaeoclimatology, Palaeoecology 37: 
MT N2. 

Pilsbry, H. A. 1921. A review of W. M. Gabb’s Tertiary Mollusca of Santo Domingo.—Proceedings 
of the Philadelphia Academy of Natural Sciences 73(2):305—428, pls. 16-47. 

Weisbord, N. E. 1962. Late Cenozoic gastropods from northern Venezuela.—Bulletins of American 
Paleontology 42(193): 1-672, 48 pls. 

Woodring, W. P. 1928. Miocene mollusks from Bowden Jamaica. Part 2, Gastropods and discussion 
of results.—Carnegie Institution of Washington, No. 385: vii + 564 pp., 39 pls. 

——. 1964. Geology and paleontology of Canal Zone and adjoining parts of Panama. Description 
of Tertiary mollusks (Gastropods: Columbellidae to Volutidae)—U.S. Geological Survey 
Professional Paper 306-C:241—297, pls. 39-47. 


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


PROC. BIOL. SOC. WASH. 
96(3), 1983, pp. 355-364 


CARENZIA, A NEW GENUS OF SEGUENZIACEA 
(GASTROPODA: PROSOBRANCHIA) WITH THE 
DESCRIPTION OF A NEW SPECIES 


James F. Quinn, Jr. 


Abstract.—Carenzia is proposed for three species of seguenziacean gastropods, 
with Seguenzia carinata Jeffreys, 1877, as type-species. Carenzia carinata and 
C. trispinosa (Watson, 1879) are redescribed and illustrated with photomacro- 
graphs and scanning electron micrographs; C. inermis is described as new and 
illustrated with photomacrographs. Carenzia carinata is known from both sides 
of the North Atlantic Ocean in depths of 1000-2000 m; C. trispinosa occurs in 
similar depths but is restricted to the western side of the North Atlantic; C. 
inermis 1s reported from off Oregon in depths of about 2000 m. 


Several species-groups within the genus Seguenzia Jeffreys, 1876, which were 
informally recognized by Quinn (in press), should be accorded generic rank. 
Diagnoses of these new genera were to be published as part of a systematic review 
of the western Atlantic species of Seguenziacea (Quinn, in preparation). How- 
ever, in a monograph of New Zealand seguenziaceans Marshall (in press) is pro- 
viding diagnoses for at least two of these genera, leaving only the Seguenzia 
carinata group to be diagnosed among western Atlantic Seguenzia. Publication 
of this paper prior to a full review is prompted by discovery of a new species of 
the S. carinata group from off Oregon which is to be included in a monograph 
of northeastern Pacific archaeogastropods (McLean, personal communication). 
Carenzia, new genus, is herewith diagnosed, and the two western Atlantic species, 
C. carinata (Jeffreys, 1877) and C. trispinosa (Watson, 1879), are redescribed 
and illustrated. Carenzia inermis, new species, is described from the northeastern 
Pacific. Two New Zealand species of Carenzia will be described by Marshall (in 
press). 

Institutional abbreviations used in this paper are: BM(NH), British Museum 
(Natural History); LACM, Los Angeles County Museum of Natural History; 
UMML, Rosenstiel School of Marine and Atmospheric Science (RSMAS), Uni- 
versity of Miami; USNM, U.S. National Museum of Natural History. 


Carenzia, new genus 
Seguenzia.—Auct. (partim). 


Type-species.—Seguenzia carinata Jeffreys, 1877; herein designated. 

Diagnosis.—Shell small, trochoid, peripherally carinate, nacreous under a por- 
celaneous layer; spire evenly conical to turreted, with mid-whorl carina at least 
on early whorls; base convex, usually with deep, wide umbilicus, occasionally 
with spiral threads; aperture rhomboidal; outer lip with shallow, V-shaped sinus 
at suture, and another in peripheral part of base; columella strongly arched, with 
or without distinct tooth; protoconch smooth or with spiral ridges, terminal varix 
low; animal, operculum and radula as yet undescribed. 


356 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Remarks.—Carenzia differs from Seguenzia in the more conical shape of the 
shell, lack of strong axial and spiral sculpture on the later whorls, V-shaped anal 
sinus, and lack of a distinct sinus in the periphery. From the genus which includes 
Fluxina discula Dall, 1889 (to be described by Marshall, in press), Carenzia 
differs in the much more elevated shell, prominent protoconch, and presence of 
a mid-whorl carination in addition to the peripheral carina. Although animals of 
this genus are undescribed, Bouchet and Warén (personal communication) have 
material of C. carinata (Jeffreys, 1877) from which they will prepare an anatom- 
ical discussion of the species. Marshall (in press) will present illustrations of the 
radula of at least one of his new species. Carenzia is known from the Atlantic 
and northeastern Pacific oceans, and New Zealand waters (Marshall, in press). 

Etymology.—From the Latin carina, a keel, and Seguenzia; gender feminine. 


Carenzia carinata (Jeffreys, 1877) 
Figs. 1-7 


Seguenzia carinata Jeffreys, 1876:201 (nomen nudum); 1877:320; 1879:606; 1885: 
43, pl. 5, figs. 3, 3a.—Kobelt, 1878:163; 1888:256.—Watson, 1879:590; 1886: 
108, pl. 7, fig. 2.—Nobre, 1884:50; 1932:182; 1936:130; 1938—40:339, 341.— 
Tryon, 1887:47, pl. 8, figs. 81-83.—Dall, 1889a:383; 1889b: 142.—Dautzenberg, 
1889:55; 1927:144.—Dautzenberg and Fischer, 1897:162; 1906:54.—Locard, 
1898:68.—Johnson, 1934:74.—? Clarke, 1961:350, pl. 4, fig. 6; 1962:11 (partim 
?).—Nordsieck, 1968:20, pl. 4, fig. 09.73.—Abbott, 1974:38.—Laubier and 
Bouchet, 1976:470—472.—Warén, 1980:15. 

Seguenzia carinata var. attenuata Locard, 1898:68. 


Material examined.—1 specimen, USNM 181657 (lectotype, herein designat- 
ed); 1 specimen, USNM 821198 (paralectotype); Valorous station 13, 56°01'N, 
34°42'W, 1263 m; 20 August 1875.—1 specimen, USNM 181658 (paralectotype); 
Porcupine station 22, 56°08'N, 13°34’W, 2311 m; July 1870.—1 specimen, USNM 
181659; Talisman station, off Azores, 1246 m.—1 specimen, BM(NH) 
1887.2.9.374a; Chailenger station 73, 38°30'N, 31°14’W, 1829 m; 30 June 1873.— 
1 specimen, BM(NH) 1887.2.9.375a; Challenger station 78, 37°26'N, 25°13'W, 
1829 m; 10 July 1873.—3 specimens, BM(NH) 1887.2.9.376-379; Challenger sta- 
tion 85, 28°42'N, 18°06'W, 2059 m; 19 July 1873.—6 specimens, UMML 30.8021; 
Gerda station G-1111, 23°51.9'N, 80°42.7’W, 1080-1089 m; 10’ otter trawl; 30 
April 1969.—3 specimens, UMML 30.8020; Gerda station G-1106, 24°02’N, 
81°30'W, 1706-1723 m; 10’ otter trawl; 29 April 1969.—8 specimens, USNM 94307; 
Blake station 34, 23°52'N, 88°56’W, 732-1098 m.—1 specimen, UMML 30.8288; 
John Elliott Pillsbury station P-604, 18°58’N, 87°28'W, 970—988 m; box dredge; 17 
March 1968.—4 specimens, BM(NH) 1887.2.9.380; Challenger station 120, 8°37'S, 
34°28'W, 1115 m; 9 September 1873. 

Description.—Shell small, conical, whorls about 5, weakly inflated, strongly 
carinate peripherally, polished, white, iridescent under porcelaneous layer. Pro- 
toconch small, about 275 wm in diameter, prominent, glassy, with minute spiral 
ridges, set off from teleoconch by fine growth discontinuity visible only under 
high magnification. Spire whorls carinated by sharp spiral angulation at or slightly 
above mid-whorl, flat above, slightly concave below angulation, smooth, some- 
times with obscure, slightly flexuous, axial lirae near the suture lines; suture 


VOLUME 96, NUMBER 3 


oma: ',.-. : ; 
. > 


Figs. 1-7. Carenzia carinata. 1, Lectotype, USNM 181657, height 2.7 mm, width 4.4 mm; 2, 
Same, basal view; 3, SEM micrograph of specimen from Gerda station G-1111, UMML 30.8021, 
apertural view, 13.4; 4, Same, close-up of whorl surface, growth lines trace outline of anal sinus, 
335x,s = suture line; 5, Same, oblique basal view, 67; 6, same, oblique view of protoconch, 134x; 
7, Specimen from Blake station 34, USNM 94307, vertical view of protoconch, 268 x. 


indistinct. Periphery marked by strong, obscurely crenulate carina, visible only 
on last whorl. Base smooth, convex to flat proximally, concave under periphery. 
Umbilicus wide, approximately 25% width of shell, bounded peripherally by spiral 
thread; ridge behind columella not developed. Aperture rhomboidal; outer lip 


358 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


bisinuate; sinuses broadly V-shaped, with rounded apices, shallow; anal sinus on 
whorl shoulder, apex just above mid-whorl carina; basal sinus in outer 4 of basal 
lip, edge very weakly flared; columella arcuate, terminating in weak, obtuse den- 
ticle. Animal undescribed. 

Measurements.—Lectotype: 2.7 mm high, 4.4 mm wide, 5+ whorls. 

Type-locality.—Valorous station 13, 56°01'N, 34°42'W, 1263 m (herein desig- 
nated). 

Remarks.—The simple, unornamented or obscurely crenulated mid-whorl cari- 
na, lack of subsutural carina and spiral threads on the base, smaller protoconch 
with spiral ridges, and weaker columellar tooth distinguish C. carinata from C. 
trispinosa (Watson, 1879), the only other species of Carenzia known from the 
North Atlantic Ocean. From C. inermis, new species, C. carinata differs in being 
smaller, having a more conical shell outline, stronger mid-whorl and peripheral 
carinae, flatter base, much wider umbilicus, and distinct columellar tooth. The 
mid-whorl carina of C. carinata is rather variable: always sharp and strong on 
the early teleoconch whorls, but in many specimens becoming weak, or even 
disappearing on later whorls. This latter condition prompted Locard (1898) to 
establish the varietal name attenuata; however, both forms may be found in the 
same lot of specimens, so no distinction of morphotypes is warranted. 

The occurrence of C. carinata is somewhat more widespread than would be 
expected from experience with species in other seguenziacean genera. Carenzia 
carinata is amphi-Atlantic in distribution, extending from the northern end of the 
Mid-Atlantic Ridge (56°01’N) to the Canary Islands (28°42'N) in the east, and to 
off the northeastern bulge of Brazil (8°37’S) in the west. Most other seguenzia- 
ceans in the Atlantic Ocean are more provincial in distribution, occurring on only 
one side of the ocean, often in or along the edges of only one or two basins 
(unpublished data). A record for the southeastern Altantic (Clarke 1961) is of 
questionable validity. As noted by Laubier and Bouchet (1976), the photographed 
specimen does not appear to be C. carinata. Since I have not examined the 
specimens, I am here including Clarke’s report in the synonymy with a query. 
Carenzia carinata 1s known principally from depths of 1000-2000 m, but has been 
recorded from as shallow as 732 m and as deep as 2311 m. Although rare in 
collections from the western Atlantic, C. carinata is rather common in depths of 
about 2000 m in the Bay of Biscay (Laubier and Bouchet 1976; Bouchet, personal 
communication). 

As noted by Warén (1980), the name carinata dates from 1877, not 1876. Jef- 
freys’ (1876) listing included no figure, and his description consisted only of the 
phrase “‘base perforated or umbilicated,’’ certainly not adequate to make the 
name available under the current rules governing zoological nomenclature. 


Carenzia trispinosa (Watson, 1879) 
Figs. 8—12 


Seguenzia trispinosa Watson, 1879:591; 1886:110, pl. 7, fig. 4.—Tryon, 1887:47, 
pl. 8, figs. 79, 80.—Dall, 1889a:268; 1889b:142; 1890:335.—Maury, 1922:90.— 
Johnson, 1934:74.—Pulley, 1952:170.—Clarke, 1962:11.—Abbott, 1974:38. 


Material examined.—1 specimen, USNM 94912; Albatross station, south of 
Cape Hatteras.—1 specimen, UMML 30.8289; Columbus Iselin station CI-356, 


VOLUME 96, NUMBER 3 359 


Figs. 8-12. Carenzia trispinosa. 8, Lectotype, BM(NH) 1887.2.9.381, height 3.35 mm, width 3.9 
mm; 9, Same, basal view; 10, SEM micrograph of specimen from Albatross station 2751, USNM 
614078, apertural view, 13.4; 11, Same, vertical view of protoconch, 134; 12, same, oblique basal 
view, 13.4x. 


24°21.8'N, 77°24.8'W, 1561-1547 m; 41’ otter trawl; 20 August 1975.—3 speci- 
mens, UMML 30.8016; Gerda station G-368, 24°03'N, 81°10’W, 961-1016 m; 16’ 
otter trawl; 15 September 1964.—1 specimen, UMML 30.8292; Gerda station 
G-370, 23°54'N, 81°19’W, 1281 m; 16’ otter trawl; 16 September 1964.—4 speci- 
mens, UMML 30.7745; Gerda station G-964, 23°46’N, 81°51’W, 1390-1414 m; 
10’ otter trawl; 1 February 1968.—S specimens, UMML 30.7758; Gerda station 
G-965, 23°45'N, 81°49’W, 1394-1399 m; 10’ otter trawl; 1 February 1968.—2 spec- 
imens, UMML 30.8023; Gerda station G-1112, 23°44’N, 81°14’W, 2276-2360 m:; 
10’ otter trawl; 30 April 1969.—1 specimen, UMML 30.8015; Gerda station G-214, 
23°43'’N, 82°49’'W, 1354-1620 m; 6’ Isaacs-Kidd midwater trawl; 20 January 
1964.—2 specimens, UMML 30.8290; Gerda station G-960, 23°30'’N, 82°26’W, 
1692-1697 m; 10’ otter trawl; 31 January 1968.—6 specimens, UMML 30.8017; 
Gerda station G-959, 23°25'N, 82°35’W, 1830 m; 10’ otter trawl; 31 January 1968.— 
1 specimen, USNM 821199; Blake station 34, 23°52'N, 88°56'W, 732-1098 m.— 


360 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


6 specimens, USNM 94306; Blake station, Yucatan Strait.—5 specimens, USNM 
96575; 4 specimens, USNM 97111; 28 specimens, USNM 330745; Albatross sta- 
tion 2751, 16°54’N, 63°12’W, 1257 m; large beam trawl; 28 November 1887.—1 
specimen, A/batross station 2117, 15°24’40"N, 63°31'30"W, 1250 m; large beam 
trawl; 27 January 1887.—23 specimens, USNM 96603; 1 specimen, USNM 406699; 
Albatross station 2760, 12°07'N, 37°17’W, 1865 m; large beam trawl; 18 December 
1887.—49 specimens, USNM 96878; Albatross station 2754, 11°40’N, 58°33’W, 
1609 m; large beam trawl; 5 December 1887.—1 specimen, UMML 30.8291; John 
Elliott Pillsbury station P-754, 11°36.9'N, 68°42.0'W, 684-1574 m; 10’ otter trawl; 
26 July 1968.—1 specimen, BM(NH) 1887.2.9.381 (lectotype, herein designated); 
3 specimens, BM(NH) 1887.2.9.382—384 (paralectotypes); Challenger station 120, 
8°37'S, 34°28’W, 1115 m; 9 September 1873.—44 specimens, USNM 330854; Al- 
batross station 2764, 36°42'S, 56°23'W, 21 m; large beam trawl; 12 January 1888. 

Description.—Shell small, conical, polished, white, iridescent under porcela- 
neous layer; whorls 6, strongly shouldered near suture, weakly to strongly cari- 
nate at mid-whorl, strongly carinate at periphery. Protoconch rather large, about 
380 wm in diameter, glassy, prominent. First 2 teleoconch whorls with sharp mid- 
whorl angulation which may become obsolete or strongly tuberculate on later 
whorls. All whorls subsequent to second with strong subsutural carina forming 
narrow, horizontal subsutural shelf; carina usually strongly tuberculate, often 
with fine spiral thread running across tops of tubercles. Periphery marked by 
strong, sharp, obscurely crenulate to tuberculate carina, visible only on last whorl. 
Base with narrow concave zone peripherally and broad convex inner area sepa- 
rated by weak, sharp spiral angulation or thread; inner area smooth and polished, 
with fine spiral striae, or with distinct spiral cords. Umbilicus wide, approximately 
25% width of shell, bounded peripherally by spiral cord and shallow groove; walls 
convex above and below strong spiral groove corresponding to columellar tooth. 
Aperture rhomboidal; outer lip bisinuate; sinuses broadly V-shaped, with rounded 
apices, shallow; anal sinus on whorl shoulder, apex about midway between sub- 
sutural and mid-whorl carinae; basal sinus slightly peripheral to middle of basal 
lip; columella strongly arcuate, terminating in rather strong, narrow, blunt tooth. 
Animal unknown. 

Measurements.—Lectotype: 3.35 mm high (measured from tip of protoconch 
to tip of columellar tooth), 3.9 mm wide, 512 whorls. Largest specimen: 4.6 mm 
high (overall), 4.8 mm wide, 6/2 whorls. 

Type-locality.—Challenger station 120, 8°37'S, 34°28’W, 1115 m. 

Remarks.—This species is the most highly sculptured of any described species 
of Carenzia. The tuberculate subsutural, mid-whorl and peripheral carinae, and 
striate base immediately distinguish C. trispinosa from C. carinata and C. iner- 
mis. As in C. carinata, C. trispinosa has a variable expression of sculptural 
elements: the tubercles on the carinae may be strong, sharp and closely spaced, 
or rather weak, blunt and widely spaced; the mid-whorl carina may become very 
reduced or absent after the first 2 or 3 teleoconch whorls; the base may be 
completely smooth, have fine impressed striae or distinct cords. All variations 
may be found in a lot with numerous specimens. 

Carenzia trispinosa has been reported only from the western Atlantic Ocean 
from off North Carolina south to off Argentina (36°42'S). Bathymetric occurrence 
of C. trispinosa is similar to that of C. carinata, having been reported from 684— 


VOLUME 96, NUMBER 3 361 


Figs. 13-16. Carenzia inermis. 13, Holotype, LACM 1806, height 6.8 mm, width 6.4 mm; 14, 
Same, basal view; 15, Paratype, USNM 784742, height 6.3 mm, width 6.3 mm; 16, Same, basal view. 


2360 m, principally between 1000 and 2000 m. The depth record of 21 m (Albatross 
station 2764, off Rio de la Plata, Argentina) is certainly a mistake. Calliotropis 
actinophora (Dall, 1890) was also collected at this station, and, like C. trispinosa, 
rarely occurs shallower than 1000 m (Quinn 1979). Since the other stations off 
the Rio de la Plata are in depths comparable to 2764, the specimens were probably 
mislabelled as to station number. Two stations may be possibilities: A/batross 
2763 (24°17'S, 42°48'30’W, off Rio de Janeiro, Brasil, 1228 m) and Albatross 2754 
(11°40’/N, 58°33’W, SE of Barbados, 1610 m). Both stations are within the normal 
depth range of C. frispinosa and simple transcription errors could change either 
to 2764. Albatross 2754 is more likely the correct station since it was near other 
stations from which C. ftrispinosa had been collected. The 30° latitudinal gap 
between the most southern undisputed localities of this species reflects the woeful 
lack of collecting effort along the northeastern coast of South America. 


362 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Carenzia inermis, new species 
Figs. 13-16 


Material examined.—1 specimen, LACM 1806; R/V Cayuse station BmT-331, 
44°33.0'N, 128°20.2’W, 2820 m; 3 November 1973 (holotype).—1 specimen, USNM 
784742; R/V Yaquima station OTB-186, 44°39.4’N, 125°36.3’W, 2800 m; 26 July 
1967 (paratype). 

Description.—Shell small, conical, whorls 5+, inflated, weakly carinate, iri- 
descent under porcelaneous layer. Protoconch and one or more teleoconch whorls 
missing from both specimens. Early remaining whorls with faint, regularly spaced, 
flexuous axial plicae above and below suture; mid-whorl area smooth except for 
obscure spiral angulation visible only with oblique lighting and high magnification; 
later whorls almost smooth, with only collabral growth lines and evanescent spiral 
threads. Periphery weakly carinated by single spiral thread, slightly overhanging 
succeeding whorl, giving slightly channeled look to suture; last “4 whorl without 
peripheral thread, slightly flattened between anal sinus and periphery. Base con- 
vex, well-rounded, smooth except for two or three obscure spiral threads near 
peripheral thread, collabral growth lines, and some spiral threads similar to those 
above periphery. Umbilicus narrow, edge defined by strong, rounded ridge, in- 
ternally constricted by broad spiral ridge, with narrow channel formed between 
circumumbilical and internal ridges. Aperture roughly quadrate; outer lip thin, bi- 
sinuate; sinuses (reconstructed from growth lines) shallow, broadly V-shaped, with 
rounded apices; anal sinus on whorl shoulder; basal sinus just peripheral to middle 
of base; columella broadly arcuate, without tooth, melding smoothly with outer 
lip, buttressed by intraumbilical ridge. Animal unknown. 

Measurements.—Holotype: 6.8 mm high, 6.4 mm wide, 5 whorls. Paratype: 
6.3 mm high, 6.3 mm wide, 4 whorls. Both specimens missing apex. 

Type-locality.—R/V Cayuse station BmT-331, off Oregon, 44°33.0’N, 
128°20.2'W, 2820 m. 

Remarks.—The relatively large, smooth shell with very weak peripheral carina 
and narrow umbilicus readily distinguishes C. inermis from C. carinata and C. 
trispinosa. Carenzia inermis 1s known only from the two types, so no discussion 
of variation, areal distribution, or bathymetry is possible at this time. 


Acknowledgments 


James H. McLean (LACM) brought the new species to my attention and kindly 
allowed me to include it in this report. In addition, he has provided a great deal 
of information without which our current understanding of the Seguenziacea would 
be considerably diminished. Bruce A. Marshall (National Museum of New Zea- 
land) provided photographs of several of his new species and generously deferred 
description of Carenzia to me. I am grateful to the following for the opportunity 
to examine specimens in their care: Joseph Rosewater and Richard S. Houbrick, 
USNM; Kathie Way, BM(NH); and Gilbert L. Voss, RSMAS. Some of the spec- 
imens examined for this report were collected during the National Geographic- 
University of Miami Deep-Sea Expeditions. W. G. Lyons, D. Crewz, and J. 
Darovec (Florida Department of Natural Resources) read and commented on this 
paper. Lana Tester (Florida Department of Natural Resources) supplied the SEM 


VOLUME 96, NUMBER 3 363 


micrographs. Sally D. Kaicher provided photomacrographs of all type material. 
This report constitutes a scientific contribution from the University of Miami. 


Literature Cited 


Abbott, R. T. 1974. American seashells. 2nd Ed.—Van Nostrand Reinhold, New York, 663 pp. 

Clarke, A. H. 1961. Abyssal mollusks from the South Atlantic Ocean.—Bulletin of the Museum of 
Comparative Zoology 125(12):345—387. 

——. 1962. Annotated list and bibliography of the abyssal marine molluscs of the world.—Na- 
tional Museum of Canada, Bulletin 181:vi + 114 pp. 

Dall, W. H. 1889a. Report on the Mollusca. Part II. Gastropoda and Scaphopoda. Reports on the 
results of dredging . . . in the Gulf of Mexico (1877-78) and in the Caribbean Sea (1879-80), 
by the U.S. Coast Survey steamer “Blake” . . . . —Bulletin of the Museum of Comparative 
Zoology 18:1—492. 

——. 1889b. A preliminary catalogue of the shell-bearing marine mollusks and brachiopods of 
the southeastern coast of the United States —Bulletin of the United States National Museum 
372 1=22 1. 

——. 1890. Preliminary report on the collection of Mollusca and Brachiopoda obtained in 1887— 
88. Scientific results of explorations by the U.S. Fish Commission steamer “‘Albatross.”’ No. 
VII.—Proceedings of the United States National Museum 12:219-362. 

Dautzenberg, P. 1889. Contribution a la faune malacologique des Iles Agores.—Résultats des Cam- 
pagnes Scientifiques accomplies sur son yacht par le Prince Albert I*, Prince de Monaco 1:1- 
112. 

——. 1927. Mollusques provenant des campagnes scientifiques du Prince Albert I* de Monaco 

dans |’Océan Atlantique et dans le Golfe de Gascogne.—Résultats des Campagnes Scientifiques 

accomplies sur son yacht par le Prince Albert I*, Prince de Monaco 72: 1—400. 

, and H. Fischer. 1897. Campagnes scientifiques de S. A. le Prince Albert I* de Monaco. 

Dragages effectués par l’Hirondelle et par la Princesse Alice, 1888—1896.—Mémoires de la 

Société Zoologique de France 10:139-234. 

, and 1906. Mollusques provenant des dragages effectués a |’ouest de |’ Afrique 

pendant les campagnes scientifiques de S. A. S. le Prince de Monaco.—Résultats des Cam- 

pagnes Scientifiques accomplies sur son yacht par le Prince Albert I*', Prince de Monaco 32: 

1-125. 

Jeffreys, J. G. 1876. Preliminary report of the biological results of a cruise in H.M.S. ‘Valorous’ 
to Davis Strait in 1875.—Proceedings of the Royal Society of London 25:177—230. 

——. 1877. New and peculiar Mollusca of the Eulimidae and other families of Gastropoda, as 
well as of the Pteropoda, procured in the ‘Valorous’ Expedition.—Annals and Magazine of 
Natural History (4)19:317-339. 

———. 1879. Notes as to the position of the genus Seguenzia among the Gastropoda.—Journal of 
the Linnean Society of London, Zoology 14:605—606. 

—. 1885. On the Mollusca procured during the ‘Lightning’ and ‘Porcupine’ Expeditions, 1868— 
70. (Part 9).—Proceedings of the Zoological Society of London for 1885:27-63. 

Johnson, C. W. 1934. List of marine Mollusca of the Atlantic coast from Laborador to Texas.— 
Proceedings of the Boston Society of Natural History 40(1): 1-204. 

Kobelt, W. 1878. Illustrirtes Conchylienbuch. Nurnberg, xvi + 331 pp. 

1886-1888. Prodromus faunae molluscorum testaceorum maria Europaea inhabitantium.— 

Nurnberg, iv + 550 pp. 

Laubier, L., and P. Bouchet. 1976. Un nouveau copépode parasite de la cavité palléale d'un gas- 
téropode bathyal dans le Golfe de Gascogne Myzotheridion seguenziue gen. sp. nov.—Archives 
Zoologie Expérimentale et Général 117(4):469-484. 

Locard, A. 1898. Expéditions scientifiques du “‘Travailleur’’ et du “‘Talisman’’ pendant les années 
1880, 1881, 1882, 1883. Mollusques testacés. II.—Paris, 515 pp. 

Marshall, B. A. In press. Recent and Tertiary Seguenziidae (Mollusca: Gastropoda) from the New 
Zealand region.—Records of the National Museum of New Zealand. 

Maury, C. J. 1922. Recent Mollusca of the Gulf of Mexico and Pleistocene and Pliocene species 
from the Gulf States. Part 2: Scaphopoda, Gastropoda, Amphineura, Cephalopoda.— Bulletins 
of American Paleontology 9(38):34—142. 


364 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Nobre, A. 1884. Moluscos marinhos do Noroeste de Portugal. Porto. 

1932. Moluscos marinhos de Portugal. Vol. 1.—Porto, 466 pp. 

—. 1936. Moluscos marinhos de Portugal. Vol. 2.—Porto, 381 pp. 

. 1938-1940. Fauna malacologica de Portugal: I. Moluscos marinhos e das aguas salobras.— 

Porto, xxxii + 807 pp. 

Nordsieck, F. 1968. Die Europaéischen Meeres—Gehauseschnecken (Prosobranchia) vom Eismeer 
bis Kapverden und Mittelmeer.—Gustav Fischer, Stuttgart, viii + 273 pp. 

Pulley, T. E. 1952. An illustrated check list of the marine mollusks of Texas.—Texas Journal of 
Science 2:167-199. 

Quinn, J. F., Jr. 1979. Biological results of the University of Miami Deep-Sea Expeditions. 130. 

The systematics and zoogeography of the gastropod family Trochidae collected in the Straits 

of Florida and its approaches.—Malacologia 19(1):1—62. 

. Inpress. A revision of the Seguenziacea Verrill, 1884 (Gastropoda:Prosobranchia). I. Sum- 

mary and evaluation of the superfamily.—Proceedings of the Biological Society of Washington. 

In preparation. A revision of the Seguenziacea Verrill, 1884 (Gastropoda: Prosobranchia). 

II. Systematics of the western Atlantic species. 

Tryon, G. W. 1887. Manual of Conchology. Vol. 9. Solariidae, lanthinidae, Trichotropidae, Sca- 
lariidae, Cerithiidae, Rissoidae, Littorinidae.—Philadelphia, 488 pp. 

Warén, A. 1980. Marine Mollusca described by John Gwyn Jeffreys, with the location of the type 
material.—Conchological Society of Great Britain and Ireland, Special Publication 1:1—60. 

Watson, R. B. 1879. Mollusca of H.M.S. Challenger Expedition. III. Trochidae, viz. the genera 
Seguenzia, Basilissa, Gaza and Bembix.—Journal of the Linnean Society of London 14:586— 
605. 

—. 1886. Report on the Scaphopoda and Gasteropoda collected by H.M.S. Challenger during 
the Years 1873-76.—Report on the Scientific Results of the Voyage of H.M.S. Challenger, 
1873-1876, Zoology 15:1—680. 


Florida Department of Natural Resources Marine Research Laboratory, 100 
Eighth Avenue SE, St. Petersburg, Florida 33701. 


PROC. BIOL. SOC. WASH. 
96(3), 1983, pp. 365-369 


THE LYCOTEUTHID GENUS OREGONIATEUTHIS VOSS, 
1956, A SYNONYM OF LYCOTEUTHIS PFEFFER, 1900 
(CEPHALOPODA: TEUTHOIDEA) 


Ronald B. Toll 


Abstract.—Oregoniateuthis Voss, 1956 is placed in the synonymy of Lycoteu- 
this Pfeffer, 1900. Based on new records, Oregoniateuthis springeri Voss, 1956 
is shown to be conspecific with Lycoteuthis diadema (Chun, 1900) and is placed 
in synonymy. The type of Lycoteuthis lorigera (Steenstrup, 1875) is shown to be 
a male. Sexual dimorphism in the family Lycoteuthidae is discussed. 


Voss (1962) erected a new genus and species, Oregoniateuthis springeri, for a 
single male specimen of a small lycoteuthid squid taken from the stomach of a 
shark from the Gulf of Mexico. Voss (1962) expanded the description of O. 
springeri based on the holotype (USNM 575090) and a second trawl-collected 
male (UMML 31.376) from near the type-locality. He also redescribed Onycho- 
teuthis (2?) lorigera Steenstrup, 1875, known only from the type taken from the 
stomach of a sperm whale from the South Pacific, and placed this species into 
Oregoniateuthis. Voss (1962) considered the possibility that O. springeri might 
be the male of some previously described species; however, the type of O. /o- 
rigera, reported to be a female, served to validate the genus by sharing with O. 
springeri elongate dorsolateral arms. 

Present findings based on new records include an undescribed photophore in 
female Lycoteuthis diadema and show that O. springeri, known previously only 
from males, is conspecific with L. diadema, known previously only from females. 

A reexamination of the type of O. lorigera shows it to be a male and further 
demonstrates that Oregoniateuthis can no longer be retained as a separate genus. 

The following abbreviations are used: USNM—National Museum of Natural 
History, formerly United States National Museum, Smithsonian Institution; 
UMML—Invertebrate Museum, Rosenstiel School of Marine and Atmospheric 
Sciences, University of Miami; SAM—South African Museum; ORE—M/V Or- 
egon; ORE II—M/V Oregon II; CI—R/V Columbus Iselin; SUL BAY—M/V Sil- 
ver Bay; IKMT—Issacs-Kidd midwater trawl; FT—flat trawl; ST—shrimp trawl; 
PT—pelagic trawl; MI—mantle length; GlL—gladius length; F(F)—female(s); 
M(M)—male(s). 


Lycoteuthis Pfeffer, 1900 


Lycoteuthis Pfeffer, 1900:156. 
Thamatolampus Chun, 1903:67. 
Asthenoteuthion Pfeffer, 1912:172. 
Leptodontoteuthis Robson, 1926:2. 
Oregoniateuthis Voss, 1956:120. 


Diagnosis.—Lycoteuthids with single large photophore embedded in muscu- 
lature of fin/mantle complex near apex of fins, with 7 additional serially arranged, 


366 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


rod-like photophores in elongate posterior projection of mantle of males (un- 
known in L. lorigera); dorsolateral arms elongate in males. 


Lycoteuthis diadema (Chun, 1900) 


Enoploteuthis diadema Chun, 1900:532. 

Lycoteuthis diadema Chun, 1903:569. 

Oregoniateuthis springeri Voss, 1956:120; 1962:283. 
(for a more detailed synonymy consult Voss 1962:275). 


Material examined.—Holotype of Oregoniateuthis springeri Voss, M, ML = 
80 mm, ORE 382, 29°11.30’'N, 88°07.30'W, 200 fms, 21 June 1951, 100’ FT, USNM 
575090. Other material:—1F, ML = 125 mm (approx.) (GL = 132 mm), CI-256, 
23°37'N, 77°06'W to 23°39'N, 77°07'W, 1360 m, in stomach of Synaphobranchus 
brevidorsalis, 1 Nov 1974, UMML 31.1735.—6FF, ML = 91-88 mm (GL = 96.2- 
93.5 mm), 35 miles from Port Elizabeth, South Africa, over 100 fms, in stomach 
of yellowfin tuna, 23 April 1974, S 38, SAM.—13FF, ML = 89-73 mm (GL = 
88.0-69.2 mm), ORE II 10906, 29°17'’N, 86°36’ W, 0-200 fms, 70’ ST, 8 Feb 1970, 
USNM 730614.—1F, ML = 79mm (GL = 85.3 mm), between Lamberts Bay and 
Vandeling Island, in stomach of Merluccius capensis, ?May 1974, S 599, SAM.— 
3FF, ML = 91-73 mm (GL = 85.2-71.3 mm), ORE II 10907, 28°36’N, 86°15’W, 
0-200 fms, 191’ ST, 9 Feb 1970, USNM 730615.—1F, ML = 82 mm (GL = 82.5 
mm), 25°09'N, 80°34’W, in stomach of Merluccius albidus, 19 Aug 1931, USNM 
576949.—1M, ML = 98 mm (GL = 73.8 mm), ORE II 11150, 24°17’N, 87°41'W, 
0-305 fms, 71’ ST, 14 Aug 1970, USNM 730611.—1M, ML = 93.7 mm (GL = 
70 mm approx.), ORE 3296, 28°36'N, 89°48'’W, 244-520 fms, 21 Aug 1961, UMML 
31.376.—3MM, ML = 95-91 mm (GL = 69.5-62.9 mm), ORE II 10907 (sta. data 
above), USNM 730612.—4MM, ML = 88-85 mm (GL = 69.2-62.8 mm), ORE II 
10906 (sta. data above), USNM 730613.—2MM, ML = 86-85 mm (GL = 65.8— 
64.2 mm), ORE II 11186, 27°49’N, 85°12'W, 0-250 fms, 120’ PT, 27 Aug 1970, 
USNM 730610.—2FF, ML = 56-46 mm (GL = 59.6-47.4 mm), SIL BAY 1198, 
24°11’N, 83°31’W, 200 fms, 9 June 1959, UMML 31.228.—1F, ML = 53 mm 
(GL = 54.5 mm), west and north of Cape Point, South Africa, in stomach of 
Thunnus alalunga?, 16 Mar 1962, A 30612, SAM.—IF, ML = 9 mm (GL = 9 
mm), ALBATROSS, 32°18'N, 63°30’ W, Ocean Acre 7-13-N, 0-1500m,3 mIKMT, 
8 Sept 1969, USNM 726955. 

Partial redescription.—In females the posterior tip of the mantle is drawn out 
into a short tail that is longitudinally cleft ventrally. A single large photophore 
lies in a pocket in the musculature at the apex of the fins. It is located slightly 
dorsoposteriorly to the convex surface of the conus of the gladius. The photo- 
phore is laterally compressed, convex dorsally and flat to slightly concave ven- 
trally. A layer of densely placed, large, dark reddish-brown chromatophores sur- 
rounds it. The opening in the musculature dorsal to the photophore is covered 
by translucent connective tissue that forms a dorsally directed window for the 
emission of light. 

See Voss (1962) for a more complete description of this and the following 
species. 


VOLUME 96, NUMBER 3 367 


Lycoteuthis lorigera (Steenstrup, 1875) 


Onychoteuthis (??) lorigera Steenstrup, 1875:473. 
Oregoniateuthis lorigera.—Voss, 1962:288. 


Material examined.—Holotype, M, ML = 180 mm, fraen Kaskelotmave, Syd- 
havet, Mus. Zool. Skibslaege. Moller. Zoologisk Museum, Kgbenhavn. 

Partial redescription.—The badly damaged male reproductive organs are small. 
The remains of | to 3 ruptured spermatophores were found in Needham’s sac. 
Only a single set of reproductive organs was found, located on the left side of 
the animal; however, the poor condition of the visceral complex does not preclude 
the possibility of paired organs. 

The posterior part of the gladius (conus) is now missing. 

Discussion.—Two fortuitous discoveries by the writer during investigations of 
gladius morphology, namely the proper determination of the gender of the type 
of Steenstrup’s O. Jorigera and the identification of a previously undescribed 
photophore in females of Lycoteuthis diadema, prompted a review of the generic 
relationships of Oregoniateuthis. 

Knowledge of the correct sex of the type of Lycoteuthis lorigera nullified the 
single substantial factor that previously substantiated Oregoniateuthis ; a nominal 
species diagnosed by elongate dorsolateral arms in the female, a character of 
males attributable to O. springeri. 

The photophore at the base of the short posterior mantle projection found in 
females of L. diadema appears to be homologous to the most proximal of the 8 
serially arranged photophores found within the much longer mantle projection of 
males attributable to O. springeri. No other lycoteuthids, with the possible ex- 
ception of L. /origera, have internal photophores similarly located. 

A review of capture data revealed that each of 2 sequential net hauls (ORE IJ 
10906 and 10907) contained both a large number of reproductively mature (gravid) 
females attributable to L. diadema (USNM 730614, 67FF and 730615, ISFF) and 
similarly developed (spermatophores present) males attributable to O. springeri 
(USNM 730612, 4MM and 730613, 13 MM). These capture records suggest a large 
mating aggregation. 

In 5 previous reports that indicate the gender of specimens attributed to either 
L. diadema or O. springeri (Chun 1910; Voss 1956, 1962, 1962a, 1967) all L. 
diadema are females, all O. springeri, males. Examination of the holdings of the 
USNM and the UMML also revealed only females of L. diadema and males of 
O. springeri (total of over 150 animals). 

These combined lines of evidence based on anatomical, behavioral, occurrence 
and distributional data provide ample grounds to place O. springeri in the syn- 
onymy of L. diadema. Retention of these as separate taxa would require that the 
males of one and the females of the other have yet to be reported. In light of the 
distribution and ecology of these animals and the extensive collecting that has 
been conducted, particularly in the Gulf of Mexico and western Atlantic Ocean, 
this possibility is untenable. 

High levels of sexual dimorphism are known to exist in the Lycoteuthidae based 
on Selenoteuthis scintillans, albeit not as great as now recognized in Lycoteuthis. 
Several important parallels in sex-linked character expression exist between Se- 


368 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


lenoteuthis and Lycoteuthis. Only the males of both genera possess brachial 
photophores. These are located at the arm tips in S. scintillans and along the 
aboral surface of the arms in L. diadema. The terminal mantle photophore in 
male S. scintillans is proportionally nearly twice the size of that found in con- 
specific females (compare Figs. 12b and 13a in Voss 1962). In L. diadema the 
posterior mantle (tail) photophores are more numerous in males (8) than females 
(1). Toll (1982) showed that L. diadema and S. scintillans share sex-linked 
character expression in 3 gladius indices. Females of both species have greater 
anterior gladius width and anterior vane length indices while males have a greater 
free rachis length index. 

Sexual dimorphism is well known in other teuthoid families. The males of both 
Alloteuthis and Uroteuthis (family Loliginidae) have a more elongate posterior 
projection of the mantle than do conspecific females. This is similar to the con- 
dition now recognized in Lycoteuthis diadema. Males of Berryteuthis anonychus 
(family Gonatidae) lack hooks on the arms while conspecific females bear hooks on 
the basal portions of arms I-III (Roper, Young and Voss 1969). In 6 genera of 
cranchiids only the females possess brachial end organs (N. Voss 1980). Among 
non-teuthoid cephalopods, oniy males of Sepia confusa possess a tail-like elonga- 
tion of the fins (Massy and Robson 1923). Males of S. incerta and S. burnupi 
possess specially modified arms (Roeleveld 1972). Sexual dimorphism in the class 
reaches its greatest expression in the pelagic octopods Argonauta and Trem- 
octopus in which the females reach a size in excess of 20 times that of the diminutive 
males. 

Imber (1975) suggested that O. springeri (=L. diadema) may be conspecific 
with L. /origera and tentatively placed it in synonymy. This is rejected in light 
of the significant difference in size at maturity. Imber’s use of /ongimanus as the 
correct senior synonym of /origera is seen as based on circumstantial evidence 
and is not followed here. 

Unfortunately, over a century after Steenstrup described L. /origera it remains 
known from only the type. Further evaluation of its taxonomic affinities must 
await additional material. Some records of females of L. diadema from southern 
latitudes may be pertinent to future consideration of L. lorigera. 


Acknowledgments 


Dr. J. Knudsen loaned the type of Onychoteuthis lorigera. Ms. M. Roeleveld 
provided specimens from South African waters. Mr. S. Hess assisted in the ver- 
ification of spermatophore remains. Drs. C. Roper, J. Rosewater and G. Voss 
and Ms. N. Voss read the manuscript and made many constructive suggestions. 
Mr. M. Sweeney assisted the writer during three visits to the Smithsonian Insti- 
tution, loaned material and read the manuscript. Juel Rembert typed the final 
manuscript. 

This work was supported in part by a Doctoral Dissertation Improvement Grant 
from the National Science Foundation (DEB-8012544) and a Predoctoral Appoint- 
ment to the Division of Mollusks from the Smithsonian Institution Fellowship 
Program. I gratefully acknowledge this support. 


VOLUME 96, NUMBER 3 369 


Literature Cited 


Chun, C. 1900. Aus den Tiefen des Weltmeeres.—Gustav Fischer, Jena, viii + 549 pp., 390 figs., 
46 pls. 

———. 1903. Aus den Tiefen des Weltmeeres. Zweite Auflage.—Gustav Fischer, Jena, x + 592 
pp., 482 figs., 46 pls. 

—. 1910. Die Cephalopoden. 1. Teil: Oegopsida.—Wissenschaftliche Ergebnisse VALDIVIA 
18(1): 1-401, atlas of 61 pls. | 

Imber, M. J. 1975. Lycoteuthid squids as prey of petrels in New Zealand seas.—New Zealand 
Journal of Marine and Freshwater Research 9(4):483—492. 

Massy, A. L., and G. C. Robson. 1923. On a remarkable case of sex-dimorphism in the genus 
Sepia.—Annals and Magazine of Natural History (9)12:435-442. 

Pfeffer,G. 1900. Synopsis der oegopsiden Cephalopoden.—Mitteilungen aus dem Naturhistorischen 
Museum 17 (Beiheft zum Jahrbuch der Hamburgischen Wissenschaftlichen Anstalten 17):145— 
198. 

——. 1912. Die Cephalopoden des Plankton Expedition.—Ergebnisse Plankton Expedition der 
Humboldt-Stiftung 2: 1-815, atlas of 48 pls. 

Robson, G. C. 1926. The Cephalopoda obtained by the S. S. PICKLE. Supplementary Report.— 
Report of the Fisheries and Marine Biological Survey, Union of South Africa 4(8): 1-6, 1 pl. 

Roeleveld, M. A. 1972. A review of the Sepiidae (Cephalopoda) of southern Africa.—Annals of the 
South African Museum 59(10): 193-313, 11 pls., 20 figs. 

Roper, C. F. E., R. E. Young, and G. L. Voss. 1969. An illustrated key to the families of the order 
Teuthoidea (Cephalopoda).—Smithsonian Contributions to Zoology 13, 32 pp., 16 pls. 
Steenstrup, J. 1875. Hemisepius, en ny slaegt of Sepia-blaeksprutternes familie, med bemaerknin- 
ger om Sepia-formerne 1 almindelighed.—Kongelige Danske Videnskabernes Selskabs Skrif- 

ter 5(10):465—482, 2 pls. 

Toll, R. B. 1982. The comparative morphology of the gladius in the order Teuthoidea (Mollusca: 
Cephalopoda) in relation to systematics and phylogeny.—Doctoral Dissertation, University of 
Miami, 390 pp., 41 pls., 12 figs. 

Voss, G. L. 1956. A review of the cephalopods of the Gulf of Mexico.—Bulletin of Marine Science 
of the Gulf and Caribbean 6(2):85—178. 

——. 1962. A monograph of the Cephalopoda of the North Atlantic I. The family Lycoteuthi- 
dae.—Bulletin of Marine Science of the Gulf and Caribbean 12(2):264—-305. 

——. 1962a. South African cephalopods.—Transactions of the Royal Society of South Africa 
36(4):245—272, 1 pl. 

—. 1967. Some bathypelagic cephalopods from South African waters.—Annals of the South 
African Museum 50(5):61-88, 9 pls. 

Voss, N. A. 1980. A generic revision of the Cranchiidae (Cephalopoda; Oegopsida).—Bulletin of 
Marine Science 30(3):365-412, 13 figs. 


Department of Invertebrate Zoology, National Museum of Natural History, 
Smithsonian Institution, Washington, D.C. 20560. Present Address: Department of 
Biology, University of the South, Sewanee, Tennessee 37375. 


PROC. BIOL. SOC. WASH. 
96(3), 1983, pp. 370-378 


PARANDALIA BENNEI (PILARGIDAE) AND 
SPIOPHANES LOWAIT (SPIONIDAE), NEW 
SPECIES OF POLYCHAETOUS ANNELIDS 

FROM MAZATLAN BAY, PACIFIC 
COAST OF MEXICO 


Vivianne Solis-Weiss 


Abstract.—Two new species of polychaetes from the Pacific coast of Mexico 
are described: Parandalia bennei and Spiophanes lowai. Both are from shallow 
sandy bottoms. 


During a recent survey of the benthic fauna of Mazatlan Bay, a number of 
polychaetous annelids new to science were encountered. Two of them are de- 
scribed below. 

The study area is the bay of Mazatlan, a shallow zone, between 3.5 and 25 
meters with active sediment transport and predominately sandy bottoms. 

The generic definitions follow Fauchald (1977). 

The types of the new species are deposited in the following institutions: Na- 
tional Museum of Natural History, Smithsonian Institution, Washington, D.C. 
(USNM), Allan Hancock Foundation (AHF), and Instituto de Ciencias del Mar 
y Limnologia, Universidad Nactional Autonoma de México (ICML). 


Family Pilargidae 
Parandalia Emerson and Fauchald, 1971 


The genus Parandalia was separated from Loandalia Monro, 1936, as emended 
by Hartman (1947:505) (see also Thomas 1963:29, and Pettibone 1966:195) chiefly 
on the basis of the presence of emergent notopodial spines or acicula in Paran- 
dalia and their absence on Loandalia. 


Parandalia bennei, new species 
Figs. 1A—D, 2 A-E 


Material examined.—Mazatlan Bay, Mexico, 23°11'55’N, 106°25'20”W, 22 Aug. 
1979, holotype ICML-1011; 2 paratypes, ICML 1011-1, ICML 1011-2; one para- 
type, USNM 073015, one paratype AHF-1345. 

Description.—The holotype is complete with 59 setigers. The four complete 
paratypes have 53, 59, 66 and 68 setigers respectively. 

The body is long and slender, 46 mm long and 0.9 mm wide, including the 
setae. It is anteriorly inflated and rounded in cross section; the middle and pos- 
terior part of the body is somewhat flattened dorsoventrally. The color of the 
preserved specimens is light brown to yellowish. The parapodia are all biramous. 
The body, after the first 7 setigers, becomes somewhat flattened dorsoventrally. 
The segmentation is rather indistinct in the anterior region, becoming very con- 


VOLUME 96, NUMBER 3 371 


Fig. 1. Parandalia bennei (holotype): A, Anterior end, dorsal view; B, Posterior end, dorsal view; 
C, Parapodium from right setiger 36, posterior view; D, Left parapodium from setiger 50, anterior 
view. Scales in mm. 


spicuous posteriorly. The segments are hexagonal in shape and the septal regions 
are narrowed. 

The prostomium is less than half as wide as the first setiger, with a longitudinal 
cleft.-between the bases of the palpophores. Distally on each of the well developed 
palpophores are located two short and retractile rodlike palpostyles (Fig. 1A). 
The proboscis was not everted in any of the specimens; however, one can see 
through the body wall that there are no jaws associated with the large mouth. 
The peristomium is not clearly distinguished from the prostomium and bears no 
tentacular cirri or any other structure. 

The parapodia are poorly developed on the first 7 setigers. The first parapodium 
bears only the neuropodium. The acicular spine protrudes from notopodium on 
all setigers starting at setiger 6. 

The neuropodia increase in size from setiger 7 to about 10 segments from the 
posterior end from which they decrease gradually. 

The notopodia are similar in size all along the body, each bearing a single thick 


372 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 2. Parandalia bennei (holotype): A, Notopodium showing spine and notoseta from middle 
setiger (2600); B, Posterior neuropodium showing bases of neurosetae (x 1000); C, Base of neuroseta 
from middle setiger (x 1100); D, Middle portion of same (2400); E, Distal ends of neurosetae (2200). 


Scanning electron microscope (SEM). 


VOLUME 96, NUMBER 3 373 


colorless emergent spine or aciculum which is short and pointed distally. In ad- 
dition, | or 2 simple slender capillary notosetae are present and there is no dorsal 
cirrus (Figs. 1C, D, 2A). 

The neuropodia bear simple capillary setae in numbers varying from 6 (ante- 
riorly) to 12 posteriorly (Fig. 1C, D). Where 12 setae are present, they emerge 
from the neuropodium in pairs (Fig. 2B). 

The neuropodium bears a single stout colorless aciculum which is thicker than 
the neurosetae. It reaches the distal edge of the neuropodium and may protrude 
slightly in some parapodia. The neurosetae are long and slender, sometimes show- 
ing double curvatures. The basal third of the neuroseta is smooth; the middle 
third has groups of fibers which fan out and sometimes show an irregular spiral 
pattern (Fig. 2C, D); in the distal third the fibers become parallel to the axis of 
the neuroseta (Fig. 2E). 

In the posterior region, the parapodia become more elongated with brown 
pigmented patches on the distal tips of the neuropodia (Fig. 1D). The last few 
segments decrease in size towards the pygidium and lose their hexagonal shapes. 
The posteriormost segment bears no setae. 

The pygidium has 3 anal cirri (Fig. 1B) including a pair of lateral long cirri and 
a shorter midventral one. The anal aperture is ventrally located but rather incon- 
spicuous. 

Remarks.—The different species of Parandalia closely resemble each other 
externally. Parandalia bennei is closest to P. fragilis, Hartmann-Schroder, 1959, 
referred to Loandalia fauveli by Pettibone, 1966. Based on Hartmann-Schroder’s 
description, the main differences between the two species are the following: 


—A pair of palpostyles are present in each palpophore in P. bennei and only 
a single palpostyle per palpophore in P. fragilis (and other Parandalia spp.). 

—Palpostyles are round and papilliform in P. fragilis, rodlike in P. bennei. 

—There is no anterior asetigerous segment present in P. bennei as there is in 
P. fragilis. 

—First parapodium is uniramous in P. bennei and biramous in P. fragilis. 

—The pygidium constitutes the most conspicuous way to separate P. bennei 
from other species: in P. fragilis the pygidium forms a pigmented, well de- 
veloped disc with small cirri. In P. bennei, the lateral cirri are large, the anal 
plate is reduced, never concave, and there is no pigmentation. 


Etymology.—The specific name is a free derivation of the name of my husband, 
to whom this species is dedicated. 

Distribution.—Parandalia bennei was found in shallow bottom areas in three 
different stations in Mazatlan Bay. There is a predominance of fine sands in two 
stations and coarse sands in one station. 


Family Spionidae 
Spiophanes Grube, 1860 
Spiophanes lowai, new species 
Figs. 3A—D, 4A—G, 5A—C 


Material examined.—Mazatlan Bay, Mexico, 23°11'55”N, 106°25’20"W, 9 m, 
25 May 1979 (holotype ICML-1012). Sinaloa coast south of Mazatlan, Mexico, 


374 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 3. Spiophanes lowai (holotype): A, Anterior end, dorsal view; B, Right parapodium, setiger 
1; C, Dorsal view of the ciliated ridges from middle part of body; D, enlargement of part of a dorsal 
ridge. Scales in mm. 


23°38'5"N, 106°55’6”W, 37 m, 23 August 1981 (2 paratypes, USNM 80179, 80467). 
The holotype is incomplete with 49 setigers, 15 mm long and 23 mm wide including 
the setae. Color (preserved) is white with dark brown parapodial glands in setigers 
10 to 15. 

The prostomium is subtriangular with very poorly developed frontal horns; it 
tapers posteriorly and extends to the level of setiger 1, with a small occipital 
antenna. 


VOLUME 96, NUMBER 3 375 


f 


ese E / 
NY ~ 

\ 

WN 


Ow 


0.001 


TC § 


Cine Dp ve. & G 


Fig.4. Spiophanes lowai (holotype): A, Left parapodium of setiger 8; B, Right parapodium of setiger 
43, anterior view; C, Neuropodial spine of first setiger; D, Capillary notoseta; E, Indented setae of 
setigers 8 and 9; F, Sabre setae; G, Neuropodial hook. Scales in mm. 


There are large pigmented areas on the dorsal part of the prostomium and at 
the level of the first 2 setigers (Fig. 3A). The palps are missing. The enlarged 
peristomium forms a conspicuous lateral collar. Nuchal organs are poorly devel- 
oped, appearing as thin bands extending to setiger 2. The neuropodia of the first 
setiger bear a long smooth spine, curved at right angles as is characteristic for 
Spiophanes (Figs. 3B, 4C). The first notopodial lamellae are cirriform (Fig. 3A, 
B). From setiger 2 to 4, the notopodial lamellae increase in size and become 
foliose in shape (Fig. 3A). More posteriorly, the notopodial lamellae become 
reduced, retaining a broad base and a slender cirrus (Fig. 4B). Neuropodial la- 
mellae are similar in shape and size through the first 17 setigers (Figs. 3B, 4A). 
More posteriorly, the lamellae become smaller and remain uniform in shape and 
size through the rest of the body (Fig. 4B). 

Most notopodial setae are capillary, unilimbate and smooth (Fig. 4D). The setae 
of setigers 2 to 4 are longer than those of the rest of the body. In the notopodia 


376 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 5. Spiophanes lowai: A, Frontal view of neuropodial hook (4400); B, Neuropodium of 
setiger 47, showing hooks and sabre setae (x440); C, Same showing basal parts of hooks (<940). 
Scanning electron microscope (SEM). 


of setigers 7 and 8 there are indented setae of the same size as the accompanying 
notosetae. Bacillary setae are present from setiger 5; each seta is long, slender, 
unsheathed and with frayed appearance. Thread glands are inconspicuous. 

Lower smooth neuropodial sabre setae first appear on setiger 5 and continue 
posteriorly, one per ramus. 

The dark colored parapodial glands are located in setigers 10 to 15. Neuro- 
podial hooks first appear in setiger 15, posterior to the parapodial glands, usually 
six hooks per ramus. Each hook is straight, lacks a hood and bears 4 teeth, the 
lowermost tooth being much larger than the other 3 (Figs. 4G, 5A—C). Five hooks 
are arranged in a vertical row with the sixth hook posterior to the lower hook 
and anterior to the sabre setae (Fig. 5C). 

Transverse dorsal ridges connecting the dorsal lamellae of the notopodia begin 
on setiger 17 and continue posteriorly to the end of the fragment. They are well 


VOLUME 96, NUMBER 3 SH 


developed, overlapping on the following segment, highly vascularized and ciliated 
(Fig. 3C, D). The first ridge is lower than the rest. 

Transparent and delicate genital pouches begin at setiger 15. The pygidium is 
unknown. 

The tube of the paratype is mucous, transparent, and covered with fine sand. 

Remarks.—Examination of specimens of §. kroeyeri Grube, 1860, and S. ber- 
keleyorum Pettibone, 1962, in the collection of the USNM, show that S. lowai 
can be differentiated from these two species as follows: 


1) The peristomium in §. /Jowai is broad and well developed as in S. berke- 
leyorum; it is reduced in S. kroeyeri. 

2) Nuchal organs are poorly developed in S. lowai. In S. berkeleyorum and in 
S. kroeyeri they are very well developed (Light 1978, and examination of 
type-material in USNM). 

3) Prostomium and anterior dorsum are pigmented in S$. Jowai. Pigmentation is 
absent in S. kroeyeri and S. berkeleyorum. 

4) Parapodial glands are present from setiger 6 to 12 in S. kroeyeri and 10 to 
15 in S. lowai. 

5) Inferior sabre setae are first present on setiger 5 in S. /owai and on setiger 
4in S. kroeyeri and S. berkeleyorum. 

6) Notopodial lamellae are similar in S. kroeyeri and S. berkeleyorum in that 
1 to 4 are digitiform, and 5 to 15 are low, rounded and semicircular according 
to Light (1978). Posteriorly they have wide obcordate bases and digitiform 
tips. In S. /owai the first lamella is digitiform, 2 to 4 increase in size and 
are foliose, and posteriorly each lamella has a wide base and digitiform tip. 

7) The dorsal transverse ridges are much more developed in S. /owai than in 
the other two species. 

8) The indented setae were not found in specimens of §. berkeleyorum and S. 
kroeyeri examined, including the types of S. berkeleyorum. 


However, in material of S. kroeyeri in the USNM collections, the specimens from 
British Columbia (Cat. No. 53249) agree with S. Jowai; other specimens identified 
as S. kroeyeri lack indented setae and the high overlapping dorsal ridges. 

Spiophanes kroeyeri is a cosmopolitan species reported from widely differing 
areas, and probably examination of further material will show that there are in 
fact more than one species involved in what is now called S. kroeyeri Grube. 

Etymology.—The specific name is a derivation of the name Lowa Weiss to 
whom this species is dedicated. 

Distribution.—The species was collected in shallow sandy bottoms in the bay 
of Mazatlan and the Sinaloa coast, south of Mazatlan. 


Acknowledgments 


I am deeply indebted to Dr. Kristian Fauchald for all the guidance provided in 
every step of this work, and without whom this work could not have been ac- 
complished. I wish to thank also Dr. Meredith Jones for his help during my stay 
in his laboratory. I am grateful to Biol. Yolanda H. de Uribe for the work done 
at the SEM (model JOEL, JSM35), to Drs. M. Hendrickx and A. Van der Heiden 
for inviting me to participate in this project and to the Instituto de Ciencias del 
Mar y Limnologia for providing the support to complete this work. 


378 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Literature Cited 


Emerson, R. R., and K. Fauchald. 1971. A revision of the genus Loandalia, Monro with description 
of a new genus and species of pilargid polychaete.—Bulletin of the Southern California Acad- 
emy of Science 70: 18-22. 

Fauchald, K. 1977. The polychaete worms. Definitions and keys to the orders, families and gen- 
era.—Natural History Museum of Los Angeles County Science Series 28:1—190. 

Foster, N. M. 1971. Spionidae (Polychaeta) of the Gulf of Mexico and the Caribbean Sea.—Studies 
on the fauna of Curagao and Canbbean Islands 36:1—183. 

Grube, A. E. 1860. Beschreibung neuer oder wenig bekannter Anneliden. Beitrag: Zahlreiche Gat- 
tungen.—Archiv ftir Naturgeschichte (Berlin) 26:71-118. 

Hartman, O. 1947. Polychaetous annelids. Part VIII. Pilargiidae.—Allan Hancock Pacific Expedi- 
tions 10:483-522. 

Hartmann-Schréder, G. 1959. Zur Okologie der Polychaeten des Mangrove-Estero-Gebietes von El 
Salvador.—Beitrage zur Neotropischen Fauna 1:69-183. 

Light, W. J. 1977. Spionidae (Annelida: Polychaeta) from San Francisco Bay, California: A revised 
list with nomenclatural changes, new records and comments on related species from the north- 
eastern Pacific Ocean.—Proceedings of the Biological Society of Washington 90:66—88. 

—. 1978. Spionidae (Polychaeta, Annelida) invertebrates of the San Francisco Bay estuaries 
system.—The Boxwood Press, Pacific Grove. xii + 211 pp. 

Monro, C. C. A. 1936. Polychaete worms. I].—Discovery Reports 12:59-198. 

Pettibone, M. H. 1962. New species of polychaete worms (Spionidae: Spiophanes) from the east 
and west coast of North America.—Proceedings of the Biological Scciety of Washington 75: 
77-88. 

——. 1966. Revision of the Pilargidae (Annelida: Polychaeta) including descriptions of new species 
and redescription of the pelagic Podarmus ploa, Chamberlin (Polynoidae).—Proceedings of 
the United States National Museum 118 (3525): 155-208. 

Thomas, J. 1963. Polychaetous worms from the Arabian Sea. I. A new species of the genus Loan- 
dalia, Monro.—Bulletin of the Department of Marine Biology and Oceanography University 
of Kerala 1:29-34. 


Instituto de Ciencias del Mar y Limnologia, Laboratorio de Ecologia Costera, 
Apartado Postal 70-305, Universidad Nacional Autonoma de México, México, 
D.F. 04510. 


PROC. BIOL. SOC. WASH. 
96(3), 1983, pp. 379-391 


AMPHISAMYTHA GALAPAGENSIS, A NEW SPECIES 
OF AMPHARETID POLYCHAETE FROM THE 
VICINITY OF ABYSSAL HYDROTHERMAL 
VENTS IN THE GALAPAGOS RIFT, AND 
THE ROLE OF THIS SPECIES 
IN RIFT ECOSYSTEMS! 


Robert Zottoli 


Abstract.—Amphisamytha galapagensis, a new polychaete species of the fam- 
ily Ampharetidae, is described from hydrothermal vents of the Galapagos Rift. 
The placement of A. galapagensis in the genus Amphisamytha and its relationship 
with the other two species in the genus are discussed. The developmental biology 
and ecology of A. galapagensis in hydrothermal vent ecosystems are described. 


Recently, I examined a collection of ampharetid polychaetes collected by the 
DSRV/Alvin from the Galapagos Rift, situated about 330 km northeast of the 
Galapagos Islands, and 640 km west of Ecuador. The Rift is part of the world- 
wide mid-oceanic ridge system where, in certain locales, warm water issues from 
hydrothermal vents. Crane and Ballard (1980) give a physical description of the 
Galapagos hydrothermal fields. Such areas often support dense communities of 
benthic animals, dominated by large vesicomyid bivalve molluscs or by mussels. 
Other organisms associated with vents include long, tube dwelling, vestimenti- 
feran worms, brachyuran and galatheid crabs, dense aggregations of serpulid 
polychaetes, eel-like fish, and many other less abundant forms (Galapagos Biol- 
ogy Expedition Participants 1979; RISE Project Group 1980). Free-living che- 
moautotrophic bacteria, which use energy derived from oxidation of hydrogen 
sulfide to convert carbon dioxide to organic carbon, most likely serve as the food 
base for such communities. Bacteria living in trophosomal tissue of the vesti- 
mentiferan tube worm, Riftia pachyptila Jones, most likely provide nutrients to 
their host by means of a similar chemoautotrophic process (Cavanaugh ef al. 
1981; Felbeck 1981; Felbeck et al. 1981; Jones 1981; Southward et al. 1981). 

The ampharetid Amphisamytha galapagensis, described here for the first time, 
is a detritivore. The external anatomy, larval development, and the role of this 
species in hydrothermal vent ecosystems are discussed. 


Materials and Methods 


Specimens were collected by the submersible DSRV/Alvin as follows: 1. Spec- 
imens, vacuumed from masses of mussel shells and the bases of vestimentiferan 
tubes with a ‘‘slurp gun,’ were preserved, and later examined at the surface; 2. 
Mussel clumps were removed from the bottom with a clam rake and placed in 


' Contribution No. 34 of the Galapagos Rift Biology Expedition, supported by the National Science 
Foundation. 


380 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


1mm 


Fig. 1. Amphisamytha galapagensis: A, Lateral view of entire worm. Setae start on segment 4. 
In younger specimens setae may be found on segment 3; B, Mid-thoracic uncinus, lateral view. 


an insulated bucket attached to Alvin’s basket; at the surface, mussels were rinsed 
over sieves with mesh openings of 0.297 and 0.88 mm; water left in the insulated 
bucket was filtered through the same sieves; animals retained on the sieves were 
preserved and examined at a later time; 3. Vestimentiferan tubes and galatheid 
crabs were treated in the same manner as mussel clumps. 


VOLUME 96, NUMBER 3 381 


16mm 


Fig. 2. Amphisamytha galapagensis: A, 11-setiger larva, lateral view; B, Larval uncinus, frontal 
view. 


The mechanical devices used to collect vent organisms are described by Ga- 
lapagos Biology Expedition Participants (1979). 


Systematics 


Ampharetid polychaetes, for the most part, are wide anteriorly and taper grad- 
ually towards the posterior end (Fig. 1A). The prostomium is trilobed. Segments 
1 and 2, which lie immediately behind the prostomium, are fused in most species 
and, ventrally, form the lower lip. Segment 3 is either achaetous or bears two 
lateral bundles of setae (paleae or winged capillary). Malmgren (1865-1866) and 
Fauvel (1927) recognized two segments in front of the paleal, making it the third; 
while Nilsson (1912), Hessle (1917), and Uschakov (1957) recognized one bi- 
annular segment, making the paleal segment the second. The former numbering 
system is used here. 


382 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Table 1.—Location, number of juveniles and adults, and source of Amphisamytha galapagensis 
examined in this study. 


Number of 
specimens? 
———____—— Maximum 
Alvin dive and vent area Date A J depth (m) Source 
878: Search for Clambake 11/19/79 0 1 2725 With galatheid crab 
879: Mussel Bed 1/20/79 1 2 2495 Mussel washings 
880: Mussel Bed 1/21/79 7 18 2493 Mussel washings 
883: Garden of Eden 1/25/79 0 6 2493 Slurp gun in mussel area 
884a: Garden of Eden 1/25/79 0 11 2482 Clam bucket mussel washings 
884b: Garden of Eden 1/25/79 12 3 2482 Rubble residue from mussel area 
887: Mussel Bed 2/12/79 3} 21 2488 Mussel Bed 
888a: Mussel Bed 2/13/79 6 51 2483 Galatheid washings 
888b: Mussel Bed 2/13/79 3 7 2483 Galatheid washings 
890: Rose Garden 2/15/79 0 9 2447 In jar with two galatheid crabs 
891: Garden of Eden 2/16/79 WE} 24 2488 Rock scrapings 
892: Rose Garden 2/17/79 2 9 2454 Residue from cooler 
894a: Rose Garden 2/19/79 0 36 2457 — 
894b: Rose Garden 2/19/79 1 8 2457 From amphipod trap? 
896-22: Rose Garden 2/21/79 26 WZ 2460 Instant ocean washings 
N983: Rose Garden 11/30/79 1 1 2457 Vestimentifera tube washings 
N98335: Rose Garden 11/30/79 0 1 2457 Vestimentifera tube washings 
N98336: Rose Garden 11/30/79 0 11 2457 Vestimentifera tube washings 
N983112: Rose Garden 11/30/79 14 es 2457 Vestimentifera tube washings 
N9846: Rose Garden 12/ 1/79 0 4 2451 Vestimentifera tube washings 
N9846c: Rose Garden 12/ 1/79 3 6 2451 Vestimentifera tube washings 
N9847: Rose Garden 12/ 1/79 0 2) 2451 Vestimentifera tube washings 
N989: Mussel Bed 12/ 6/79 3 8 2482 Bottle rack washings 
N99011: Rose Garden 12/ 7/79 0 1 2451 Vestimentifera tube washings 
N99013: Rose Garden 12/ 7/79 1 0 2451 Vestimentifera tube washings 
N99014: Rose Garden 12/ 7/79 0 D 2451 Vestimentifera tube washings 
N99019: Rose Garden 12/ 7/79 1 1 2451 Vestimentifera tube washings 
N99040: Rose Garden 12/ 7/79 0 1 2451 Vestimentifera tube washings 
N99041: Rose Garden 12/ 7/79 16 11 2451 Vestimentifera tube washings 
N9931: Garden of Eden 12/10/79 7 6 2518 Vestimentifera tube washings 
N9932: Garden of Eden 12/10/79 2 0 2518 Vestimentifera tube washings 
N9934: Garden of Eden 12/10/79 5) 0 2518 Vestimentifera tube washings 


Search for Clambake 00°47.92'N, 86°13.5’W 
Mussel Bed 00°47.89’N, 86°09.21'W 

Garden of Eden 00°47.69'N, 86°07.74’W 
Rose Garden 00°48.25’N, 86°13.48’W 


1 A = adult, J = juvenile. 


Amphisamytha galapagensis, new species 
Figs. 1A, B, 2A, B 


Material examined.—See Table 1. Holotype, USNM 81288; 13 Paratypes, 
USNM 81289. 

Description.—Holotype, gravid female, 9.6 mm long, 1.2 mm wide, with 30 
setigerous segments. Other specimens up to 10.2 mm long and 1.5 mm wide with 
32 setigerous segments. Sexually mature worms as small as 3.6 mm in length. 
Worms longer than this considered adults. About 300 elliptical eggs, 40-150 um 
at their widest point, in body cavity of one 9-mm-long female; 60 eggs at maximum 


VOLUME 96, NUMBER 3 383 


Table 2.—Number of smooth winged capillary setae per notopodium on “‘setigerous’’ segments of 
the 11- to 18-setiger stages of Amphisamytha galapagensis. 


Setiger stage 


Segment Setiger lla 11b 14 18a 18b 18¢ 18d 
3 1 3 2 3 2 l 0 
4 2 3 3 3 3 4 z 5 
5 3 3 4 3 4 5 4 6 
6 4 3 5 3 5 5 5 6 
7 5 3 4 4 4 5 4 6 
8 6 3 3 5 6 5) 5) 6 
9 7 3 3 3 6 4 5 6 
10 8 3 3 3 3 4 5 7 
11 9 3 2, ?) 4 4 4 7 
12 10 2, 2 2 5 4 | 5 7 
13 11 1 | 2 4 4 4 7 
14 12 1 4 4 4 7 
15 13 1 4 3 4 6 
16 14 1 4 3 4 i] 
17 15 4 yy 3 7 
18 16 2 Z 3 7 
19 17 2 3 6 

20 18 2 Zz 2 5 
Ila = 0.5 x 0.14 mm 18b = 1.5 x 0.35 mm 
11b = 0.7 x 0.18 mm 18c = 1.75 x 0.4 mm 
14 = 0.8 x 0.18 mm 18d = 8.0 x 1.2 mm 


18a = 1.05 x 0.25 mm 


diameter. Color of adult in alcohol white to pale orange. Prostomium indistinctly 
trilobed, lacking glandular ridges. About 25 smooth, ventrally grooved oral ten- 
tacles. Segments | and 2 fused, with ventral part forming lower lip. Segment 3 
lacking paleae. Minute notopodial lobes with 2—3 short winged capillary setae on 
segment 3 in juveniles and smaller adults. Each seta approximately 0.25 mm long, 
5 wm wide basally, and 6 wm wide across the blade. Four pairs of smooth bran- 
chiae about % of body length, individually inserted, across dorsal surface of 
segments 3—5. Origin of branchial pairs described in larval development section. 
Wide gap between branchial groups. Notopodial lobes bearing 6-10 smooth winged 
capillary setae from segment 4 to end of thorax. Each seta about 0.75 mm long, 
10 «wm wide basally and 13 wm wide across blade. Notopodia of segments 4 and — 
5 raised dorsally; notopodia of segment 4 smaller than others. 17 thoracic setig- 
erous segments. Neuropodial lobes (uncinigerous pinnules) each with single row 
of toothed uncini from segment 7 to end of abdomen. 14 uncinigerous thoracic 
and 12-15 uncinigerous abdominal segments. Thoracic and abdominal uncini in 
single transverse rows with, respectively, about 20 and 15 per row in larger 
specimens. Thoracic and abdominal uncini with 4 teeth in single row above round- 
ed basal prow (Fig. 1B). Larval uncini may be found in juveniles and young adults 
(Fig. 2B). No notopodial, neuropodial, or anal cirri. Notopodia absent from ab- 
dominal segments. Thickened glandular pads above uncinigerous pinnules on 
most abdominal segments. Pygidium bilobed. 

Remarks.—The genus Amphisamytha was created by Hessle (1917) to include 
ampharetids with four pairs of smooth branchiae, no setae on segment 3, no 


384 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


glandular ridges on the prostomium, 14 thoracic uncinigerous segments and no- 
topodia on abdominal segments. 

Day (1964) considered Amphisamytha Hessle (1917) and Hypaniola Annekova 
(1928) to be synonyms of Lysippides Hessle (1917). He expanded this genus to 
include those with or without setae on segment 3. Admittedly the three genera 
are closely allied; however, the complete lack of setae on segment 3 in larger 
specimens, I feel, justifies the separation of Amphisamytha from these closely 
related genera. 

Amphisamytha galapagensis is distinguished from the other two members of 
the genus, A. bioculata (Moore) and A. japonica (Hessle), by the complete lack 
of anal cirri and by the possession of thick glandular pads above uncinigerous 
pinnules in most abdominal segments. 

Etymology.—Specific name refers to geographic area where specimens were 
collected. 


Feeding 


Amphisamytha galapagensis lives in mucus-lined tubes covered by detritus 
and small chips of volcanic glass, which resemble tubes of the brackish water 
ampharetids Hypania invalida (Grube) and Hypaniola kowalewskii (Grimm) 
(Manoleli 1977). The tubes are attached to solid surfaces such as lava or clam 
shells. 

The morphology of the feeding apparatus of A. galapagensis is remarkably 
similar to that described for the majority of ampharetid species. This suggests a 
similarity in feeding behavior. Below is a description of feeding behavior, from 
the sources noted, for Asabellides oculata (Webster) and Hobsonia (Amphicteis) 
floridus (Hartman) (Fauchald and Jumars 1979; personal observations); Hypania 
invalida (Grube) (Manoleli 1975); and Melinna palmata (Grube) (Dragoli 1961). 
In general ampharetids feed by extending tentacles from the tube and over the 
sediment surface. Mucus, secreted in the ventral part of each tentacle, traps 
detritus which is carried by ciliary action along the ventral tentacular groove to 
the mouth. Manoleli (1975) noted that the tentacles are suddenly extended to 
about three-fourths of the body length from the tube onto the sediment surface; 
then, laden with detritus, they are gradually retracted towards the mouth, some- 
times independently of one another. Similar feeding behavior was noted by the 
author for A. oculata (Webster) and by Dragoli (1961) for M. palmata (Grube). 
Although most ampharetids are considered to be detritus feeders, Manoleli (1975) 
suggested that H. invalida (Grube) and H. kowalewskii (Grimm) are suspension 
feeders since phytoplankton is common in gut cavities. 

The presence of detritus and bacteria in digestive tracts of juvenile and adult 
A. galapagensis lends support to the idea that mainly they consume particulate 
matter that settles on solid surfaces; however, suspension feeding cannot be ruled 
out. The bulk of their food most likely consists of chemoautotrophic bacteria and 
fecal pellets from other organisms. 


Reproduction and Larval Development 


Sexually mature worms range from 3.6 to 10.2 mm long and from 0.75 to 1.5 
mm wide. Based on observations on other ampharetid species, eggs probably 


VOLUME 96, NUMBER 3 385 


Frequency 


0 IL 2 3 4 5 6 7 8 9) 10 ileal 
Length in mm 


Fig. 3. Frequency histogram for length of a collection of Amphisamytha galapagensis from the 
Rose Garden on Alvin dives 890, 892, 894, 896 (clear portion); 983, 894 and 990 (shaded portion). 
Refer to systematics section for dive locations. 


pass singly through nephridia and nephridiopores into the anterior part of the 
tube where they are fertilized by sperm released by males in the same fashion. 
Gametes of the ampharetid Melinna cristata (Sars) are ejected in two streams 
from the anterior end of the tube (Nyholm 1950) while Fauvel (1897) indicated 
that Ampharete grubei (Malmgren) and Amphicteis gunneri (Sars) shed eggs sin- 
gly into the water column through two nephridia on the fourth setigerous segment. 

Unfertilized eggs, between 40-150 wm maximum width, are irregularly elliptical 
with a large germinal vesicle. Developing larvae beyond the fertilized egg and 
prior to the 10-setiger stage were not found. Larval stages in this paper are named 
on the basis of the number of segments (setigers) bearing capillary setae. In an 
adult, however, setiger refers to any segment bearing setae of the capillary or 
uncinate type. 

The 11-setiger stage, illustrated in Fig. 2A, is described below. Larval ciliary 
bands (prototroch, metatrochs or telotroch) were not evident. They disappear at 
about the 6-setiger stage in H. floridus (Hartman), which has similar larval de- 
velopment (Zottoli 1974). There is a single, median, ventrally grooved and ciliated 
tentacle protruding from the mouth. In the 8-setiger stage of H. floridus (Hartman) 


386 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Table 3—Number of larval uncini per uncinigerous pinnule on “‘setigerous”’ and “‘post-setigerous”’ 
segments of the 11- to 18-setiger stages of Amphisamytha galapagensis. Larval sizes are listed in 
Table 2. 


Setiger stage 


Segment Setiger lla 11b 14 18a 18b 18c 18d 
3 1 0 0 0 0 0 0 0 
4 2 1 0 0 0 0 0 0 
5 3 1 1 0 0 0 0 0 
6 4 3 1 1 0 0 0 0 
7 5 3 1 1 5 2 7 0 
8 6 3 2 4 5) 3 7 0 
9 7 3 2 4 4 3 6 0 
10 8 D 2 3 4 3 4 0 
11 9 2 y, 3 4 2 3 0 
12 10 1 1 2, 4 2 3 0 
13 11 1 1 2) 3 3 3 0 
14 12 1 1 2 3 3 3 0 
15 13 1 1 1 3 3 3 0 
16 14 1 3 3 3 0 
17 15 1 3 2 3 0 
18 16 1 3 2 3 0 
19 17 2 3 3 0 

20 18 D, 3 3 0 

““Post- 

setiger”’ 

(abdomen) 

21 1 2 6 5 0 
aD 2 2 5 5) 0 
23 3 5 4 0 
24 4 4 4 0 
25 5 3 2; 0 
26 6 2 1 0 
27 7 1 1 0 
28 8 0 
29 9 0 
30 10 14 
31 11 10 
32 12 Oe 
33 13 6 


this tentacle is used in feeding, and detrital particles, bound in mucus secreted 
at the tentacle tip, are transported by ciliary action along the ventral groove to 
the mouth. Prostomial eyespots are lacking and fused segments | and 2 lie im- 
mediately behind prostomium. Segments 3—5 bear small notopodia, each with one 
spatulate and one smooth winged capillary seta. Spatulate setae are larval struc- 
tures formed only in notopodia of the first three setigers. They disappear in later 
stages. Similar setae were described by Wilson (1928) on the first 11 setigers of 
the terebellid polychaete Loimia medusa (Savigny). One uncinus is embedded in 
the epidermis on each side of segments 4 and 5 below the notopodia. Each uncinus 
bears about 6-10 teeth above a single tooth and rounded basal part (Fig. 2B). 
Segments 6-13 each bear a pair of notopodia bearing smooth, winged capillary 
setae (Table 2). Uncinigerous pinnules, each have larval uncini on segments 7— 


VOLUME 96, NUMBER 3 387 


Table 4.—Number of adult uncini per uncinigerous pinnule on ‘“‘setigerous’’ and “‘post-setigerous”’ 
segments of the 11- to 18-setiger stages of Amphisamytha galapagensis. Larval sizes are listed in 
Table 2. 


Setiger stage 


Segment Setiger Ila 11b 14 18a 18b 18c 18d 
3 1 0 0 0 0 0 0 0 
4 D, 0 0 0 0 0 0 0 
5 3 0 0 0 0 0 0 0 
6 4 0 0 0 0 0 0 0 
V 5 0 0 0 0 5) 1 18 
8 6 0 0 0 0 6 1 18 
9 7 0 0 0 0 4 2 22 
10 8 0 0 0 0 4 3 25 
11 9 0 0 0 0 3. 2 23 
12 10 0 0 0 0 3 3 20 
13 11 0 0 0 0 3 3 17 
14 12 0 0 2 3 20 
15 13 0 0 D ) 19 
16 14 0 0 D) D 19 
17 15 0 2 2 18 
18 16 0 3 Dp 17 
19 17 0 1 Dy, 15 

20 18 0 1 1 14 

““Post- 

setiger”’ 

(abdomen) 

21 1 0 0 15 
OD 2 0 0 14 
23 3 0 0 15 
24 4 0 0 15 
25 5 0 0 13 
26 6 0 0 14 
27 7 0 0 14 
28 8 14 
29 9 15 
30 10 2 
31 11 0 
32 12 0 
33 13 0 


13 (Fig. 2A, Table 3). First and only pair of branchiae originate from dorsal 
surface of segment 3. As growth continues the uncini disappear from segments 4 
and 5. . 

In the 14-setiger stage, the second pair of branchiae arise behind the first on 
dorsal surface of segment 4; larval uncini disappear from segment 6. 

The following changes take place between the stages above and adult worms: 
1) Smooth winged capillary setae on segment 3 are lost. This suggests that pos- 
session of setae on segment 3 is a primitive character in the family Ampharetidae. 
2) Abdominal segments increase in number to 12-15. 3) Larval spatulate setae 
are lost. 4) Uncini change in number and structure: a) There is an increase in 
number of adult uncini per uncinigerous pinnule (Fig. 1B, Table 4) coinciding 


388 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


with a decrease in number of larval uncini (Fig. 2B, Table 4). Uncini are first 
formed in the dorsal part of each pinnule. b) As new uncini are formed, older 
ones are pushed ventrally. Usually, one developing uncinus is visible in each 
abdominal uncinigerous pinnule of young worms. Uncini of the maldanid poly- 
chaetes Clymenella torquata (Leidy) and Euclymene oerstedi (Claparede) are 
formed ventrally rather than dorsally as above (Pilgrim 1977). This character may 
aid in determining general evolutionary relationships between polychaete fami- 
lies. Length of individual A. galapagensis uncini in each row increases slightly 
from ventral to dorsal supporting the idea that uncini move along the row. Pilgrim 
(1977:294) made the point that in maldanids, “‘the difference in length between 
ventral and dorsal chaetae in older worms becomes an indicator of their rate of 
progress along the row, and presumably of the growth rate of the whole body, 
the greater difference in length, the lower the growth rate.’’ As uncini of A. 
galapagensis differ slightly in length along each row, a rapid growth rate is sug- 
gested. Larval uncini are still present in the ventral part of adult posterior ab- 
dominal uncinigerous pinnules. Since newly formed uncini are of the adult type, 
the presence of larval uncini suggests that once a particular size is reached, 
growth slows. This would allow food reserves to be used for gamete production 
and development, rather than growth, thus enhancing the reproductive potential 
of the population. 5) A fourth pair of branchiae appears on dorsal surface of 
segment 5. 6) Up to 25 smooth, ventrally grooved and ciliated, oral tentacles are 
developed. 

Development of this species is similar to that of the ampharetids A. grubei 
Malmgren (Thorson 1946), H. floridus (Hartman) (Zottoli 1974), M. cristata (Sars) 
(Nyholm 1950), and Schistocomus sovjeticus Annekova (Okuda 1947). 

Environmentally predictable deep-sea communities are inhabited for the most 
part by k-selected species, characterized by low reproductive potentials, long life 
spans, and long maturation periods (Sanders and Hessler 1969; Sanders 1979). 
One might predict, therefore, that numbers and biomass of juveniles in the deep- 
sea would tend to be lower than those of adults over a given time span. This 
hypothesis is supported by data on deep-sea bivalves (Grassle and Sanders 1973), 
two isopod species (Hessler 1970), one species of tanaid (Gardiner 1975), and 
certain large bathypelagic euphausiids (Mauchline 1972). The deep-sea brittle-star 
Ophiura ljungmani (Lyman), however, is an exception to this rule (Tyler and 
Gage 1980). 

Lonsdale (1977) observed large numbers of dead mussels in certain vent areas 
suggesting that this unique environment is unstable. Instability tends to favor 
opportunistic or r-selected species (Grassle and Grassle 1974) that are character- 
ized by high reproductive rate, short maturation time, short life span, large pop- 
ulation size, high mortality rate, wide physiological tolerances and broad cos- 
mopolitan distributions (Sanders 1979). High reproductive rates allow opportunists 
to increase numbers under favorable conditions, thus enhancing their ability to 
colonize new areas, and ensure survival of the species. The deep-sea wood boring 
bivalve Xylophaga (Turner 1973), the ampharetid polychaete Decemunciger, as- 
sociated with Xylophaga (Zottoli 1982), possibly the Galapagos Rift vesicomyid 
clams (Turekian and Cochran 1981) and undescribed mussels (Rhoads ef al. 1981) 
have these characters more than many other deep-sea organisms. 

The presence of numerous (300) small eggs (150 wm maximum diameter) in the 


VOLUME 96, NUMBER 3 389 


body cavity of some female worms and the presence of numerous juveniles in 
most samples (Fig. 3) imply a high reproductive rate and suggest that A. gala- 
pagensis is relatively opportunistic. Long planktonic development associated with 
opportunism would improve the chances of worms reaching and colonizing new 
vent areas. Egg size is often suggestive of whether or not a species has planktonic 
development. Thorson (1951) suggested that polychaete species with egg diam- 
eters less than 150 um generally have long pelagic planktotrophic development. 
This generalization does not apply to most ampharetids investigated to date. H. 
floridus (Hartman) (Zottoli 1974), H. kowalewskii (Grimm) (Marinescu 1964) and 
Melinnexis arctica Annekova (Annekova 1931), which have eggs roughly the 
same width as A. galapagensis, retain developing larvae in the maternal tube 
until they are able to crawl on the bottom. Eggs of Alkamaria romijni Horst, a 
protandric hermaphrodite, either develop in the body cavity. or in the maternal 
tube (Wesenberg-Lund 1934). On the basis of its similarity to the above mentioned 
ampharetids, it is here hypothesized that A. galapagensis produces larvae that 
swim or crawl along the bottom in the area where they were released. This would 
allow continuous repopulation without relying on larvae from geographically sep- 
arated areas. As regards emigration to new sites, larvae would probably be 
swept by bottom currents to new vent areas. Lonsdale (1977) recorded a current 
speed of 18 cm/sec along the bottom in a vent area on the East Pacific Rise. Cold 
water would reduce the metabolic rate as larvae are swept away from warm vent 
areas, thereby prolonging survival time and enhancing the chances of reaching a 
new vent site. 


Acknowledgments 


Thanks are due to Dr. J. Fred Grassle, for making specimens available and to 
Dr. J. Fred Grassle, Charlene D. Long, and Dr. Meredith Jones for reviewing 
the manuscript. The study is contribution No. 34 of the Galapagos Rift Biology 
Expedition, supported by the National Science Foundation. 


Literature Cited 


Annekova, N. 1928. Uber die pontokaspischen Polychaeten. 2. Die Gattungen Hypaniola, Parhy- 
pania, Fabricia und Manajunkia.—Annuaire du Musée Zoologique de |’ Academie des Sciences 
de !URRS 30: 13-20. 

1931. Zur Polychaetenfauna von Franz-Joseph-Land (Melinnexis gen. nov. arctica sp.n.).— 
Zoologischer Anzeiger, Leipzig 95:269-272. 

Cavanaugh, C. M., S. L. Gardiner, M. L. Jones, H. W. Jannasch, and J. B. Waterbury. 1981. 
Prokaryotic cells in the hydrothermal vent tube worm Riftia pachyptila Jones: Possible che- 
moautotrophic symbionts.—Science 213:340-342. 

Crane, K., and R. D. Ballard. 1980. The Galapagos Rift at 86°W: 4. Structure and morphology of 
hydrothermal fields and their relationship to the volcanic and tectonic processes of the Rift 
Valley.—Journal of Geophysical Research 85(B3): 1443-1445. 

Day, J. H. 1964. A review of the family Ampharetidae (Polychaeta).—Annals of the South African 
Museum 48(4):97—120. 

Dragoli, A. L. 1961. Peculiar feeding habits in the Black Sea polychaete Melinna palmata Grube.— 
Doklady Akademii Nauk SSR 138(4):970—973. [In Russian. ] 

Fauchald, K., and P. A. Jumars. 1979. The diet of worms: A study of polychaete feeding guilds.— 
Oceanography Marine Biology Annual Review 17:193—284. 

Fauvel, P. 1897. Recherches sur les Ampharétiens, Annélides polychetes sédentaires. Morphologie, 


390 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Anatomie, Histologie, Physiologie.—Bulletin Scientifique de la France et de la Belgique 30: 
277-488. 
1927. Polychetes Sédentaires—Faune de France 16:1—494. 

Felbeck, H. 1981. Chemautotrophic potential of the hydrothermal vent tube worm Riftia pachyptila 

Jones (Vestimentifera).—Science 213:336—338. 

, J. J. Childress, and G. N. Somero. 1981. Calvin-Benson cycle and sulphide oxidation 

enzymes in animals from sulphide-rich habitats.—Nature 293:291—293. 

Galapagos Biology Expedition Participants. 1979. Galapagos ‘79: Initial findings of a deep-sea bi- 
ological quest.—Oceanus 22(2):2—-10. 

Gardiner, L. F. 1975. The systematics, postmarsupial development, and ecology of the deep-sea 
family Neotanaidae (Crustacea: Tanaidacea).—Smithsonian Contributions to Zoology 170:1— 
265. 

Grassle, J. F., and J. P. Grassle. 1974. Opportunistic life histories and genetic systems in marine 

benthic polychaetes.—Journal of Marine Research 32:253—284. 

, and H. L. Sanders. 1973. Life histones and the role of disturbance.—Deep-Sea Research 

20:642-659. 

Hessle, C. 1917. Zur kenntnis der terbellomorphen Polychaeten.—Zoologische Bidrag fran Uppsala 
5:39-258. 

Hessler, R. R. 1970. The Desmosomatidae (Isopoda, Asellota) of the Gay Head-Bermuda tran- 
sect.—Bulletin of the Scripps Institution of Oceanography 15:1-185. 

Jones, M. 1981. Riftia pachyptila Jones: Observations of the vestimentiferan worm from the Ga- 
lapagos Rift.—Science 213:333-336. 

Lonsdale, P. 1977. Clustering of suspension feeding macrobenthos near abyssal hydrothermal vents 
at oceanic spreading centers.—Deep-Sea Research 24:857-863. 

Malmgren, A. J. 1865-1866. Nordiska Hafs-Annulater.—Ofversigt Svenska Vetenskaps Akade- 
miens Foérhandlingar 22:181—192, 355-410. 

Manoleli, D. 1975. On the distribution, biology and origin of polychaeta from the Danube and 
Danube delta.—Travaux du Muséum d’Histoire Naturelle ‘“‘Grigore Antipa’’ 16:25—34. 

. 1977. La structure, la texture et la composition minéralogique des tubes de Hypania invalida 
et Hypaniola kowalewskii (Polychaeta, Ampharetidae) du Danube. Travaux du Muséum d’His- 
toire Naturelle “‘Gngore Antipa’’ 28:9-15. 

Marinescu, V. P. 1964. La reproduction et le développement des polychetes reliques Ponto-Caspiens 
du Danube: Hypaniola kowalewskii (Grimm) et Manayunkia caspica.—Revue Roumaine de 
Biologie, Série de Zoologie 9:87—100. 

Mauchline, J. 1972. The biology of bathypelagic organisms, especially Crustacea.—Deep-Sea Re- 
search 19:753-780. 

Nilsson, D. 1912. Beitrage zur kenntnis des nervensystems der polychaeten.—Zoologiska Bidrag 
fran Uppsala 1:85-161. 

Nyholm, K. 1950. Contributions to the life-history of the ampharetid, Melinna cristata.—Zoologiska 
Bidrag fran Uppsala 29:79-91. 

Okuda, S. 1947. On an ampharetid worm, Schistocomus sovjeticus Annekova, with some notes on 
its larval development.—Journal Faculty Science Hokkaido Imperial University (ser. 6) 9:321— 
329. 

Pilgrim, M. 1977. The functional morphology and possible taxonomic significance of the parapodia 
of the maldanid polychaetes Clymenella torquata and Euclymene oerstedi.—Journal of Mor- 
phology 152:281-302. 

Rhoads, D. C., R. A. Lutz, E. C. Revelas, and R. M. Cerrato. 1981. Growth of bivalves at deep- 
sea hydrothermal vents along the Galapagos Rift Science 214:911—-913. 

RISE Project Group. 1980. East Pacific Rise: Hot springs and geophysical experiments.—Science 


207: 1421-1444. 
Sanders, H. L. 1979. Evolutionary ecology and life-history patterns in the deep-sea.—Sarsia 64: 
7. 


, and R. R. Hessler. 1969. Ecology of the deep-sea benthos.—Science 163:1419-1424. 

Southward, A. J., E. C. Southward, P. R. Dando, G. H. Rau, H. Felbeck, and H. Fliigel. 1981. 
Bacterial symbionts and low #C/??C ratios in tissues of Pogonophora indicate unusual nutrition 
and metabolism.—Nature 293:616—620. 

Thorson, G. 1946. Reproduction and larval development of Danish marine bottom invertebrates, 


VOLUME 96, NUMBER 3 391 


with special reference to the planktonic larvae in the Sound (@resund).—Meddeleser Fra 
Kommissionen for Danmarks Fiskeri-og Havundersogelser, serie: Plankton 4(1):1—523. 

———. 1951. Zur jetzigen lage der marinen bodentierdkologie —Verhandlungen der Deutschen 
Zoologischen Gesellshaft 12(17):276—326. 

Turekian, K. K., and J. K. Cochran. 1981. Growth rate of a vesicomyid clam from the Galapagos 
spreading center.—Science 214:909-911. 

Turner, R. D. 1973. Wood-boring bivalves, opportunistic species in the deep-sea.—Science 180: 
1377-1379. 

Tyler, P. A., and J. D. Gage. 1980. Reproduction and growth of the deep-sea brittlestar Ophiura 
ljungmani (Lyman).—Oceanolica Acta 3(2):177-185. 

Uschakov, P. V. 1957. Bristleworms of the far eastern seas of USSR (Polychaeta). Opredeliteli po 
faune SSSR (Keys to the fauna of the USSR) 56:1—445. [In Russian. ] 

Wesenberg-Lund, E. 1934. A viviparous brackish-water ampharetid, Alkmaria romijni Horst from 
Ringkgbing Fyord.—Videnskabelige Meddelelser Naturhistorisk Forening 98:215—222. 

Wilson, D. P. 1928. The post-larval development of Loimia medusa Sav.—Journal of the Marine 
Biological Association of the United Kingdom 15(1):129-147. 

Zottoli, R. A. 1974. Reproduction and larval development of the ampharetid polychaete Amphicteis 
floridus.—Transactions of the American Microscopical Society 93(1):78-89. 

———. 1982. Two new genera of deep-sea worms of the family Ampharetidae and the role of one 
species in deep-sea ecosystems.—Proceedings of the Biological Society of Washington 95(1): 
48-57. 


Department of Biology, Fitchburg State College, Fitchburg, Massachusetts 
01420. 


PROC. BIOL. SOC. WASH. 
96(3), 1983, pp. 392-399 


A NEW SCALE WORM (POLYCHAETA: POLYNOIDAE) 
FROM THE HYDROTHERMAL RIFT-AREA OFF 
WESTERN MEXICO AT 21°N 


Marian H. Pettibone 


Abstract.—A unique polynoid polychaete, Lepidonotopodium fimbriatum n. 
gen., n. sp., is described from the hydrothermal vent area at 21°N off western 
Mexico and referred to a new subfamily of Polynoidae, the Lepidonotopodinae. 


During Alvin dives 1213 and 1223 in 1982 to the East Pacific Rise hydro- 
thermal vent area at 21°N in the Pacific Ocean off western Mexico, three adults 
and one young specimen of a unique polynoid polychaete were collected at 2600 
meters depth in the Clam Acres area. They were made available to me by Dr. 
Kenneth Smith of Scripps Institution of Oceanography, University of California, 
San Diego. This is OASIS Expedition Contribution number 10. 

The types are deposited in the National Museum of Natural History, Smith- 
sonian Institution (USNM). 


Family Polynoidae 
Lepidonotopodinae, new subfamily 
Lepidonotopodium, new genus 


Type-species.—Lepidonotopodium fimbriatum, new species. 

Diagnosis.—Body short, flattened, subrectangular; 28 segments (first achae- 
tous). Elytra and prominent elytrophores 11 pairs, on segments 2, 4, 5, 7, 9, 11, 
13, 15, 17, 19, 21, with dorsal cirri on posterior 7 segments. Prostomium deeply 
bilobed; median antenna with ceratophore inserted in anterior notch, with short 
style; without lateral antenna but with small frontal filaments on subtriangular 
lobes of prostomium; paired palps cylindrical, smooth, with filamentous tips; 
without eyes. First or tentacular segment fused to prostomium, not visible dor- 
sally; tentaculophores lateral to prostomium, each with pair of tentacular cirri, 
single aciculum, without setae; without facial tubercle. Second segment with first 
pair of elytra, biramous parapodia, and ventral or buccal cirri attached to basal 
parts of parapodia lateral to mouth; styles longer than following ventral cirri. 
Parapodia biramous, with notopodia shorter than neuropodia. Notopodia sub- 
conical, with projecting acicular processes and well-developed bracts encircling 
notopodia anteriorly and dorsally. Neuropodia diagonally truncate, deeply notched 
dorsally, without projecting acicular processes. Distal tips of notopodia, noto- 
podial bracts, and neuropodia fimbriated with slender papillae. Notosetae nu- 
merous, subequal in width to neurosetae, with single rows of widely spaced spines 
and blunt tips. Neurosetae numerous, with 2 rows of numerous spines along one 
side and slightly hooked blunt tips. Dorsal cirri on segments lacking elytra, with 
cylindrical cirrophores attached on posterodorsal sides of notopodia, with tapered 
styles; ventral cirri short, tapered, attached near middle of neuropodia. Dorsal 
tubercles on cirrigerous segments large, inflated. Elytrophores, dorsal tubercles 


VOLUME 96, NUMBER 3 393 


Fig. 1. Lepidonotopodium fimbriatum, holotype, USNM 80113: Dorsal view, left; ventral view, 
right. 2x. 


and their bases with numerous ciliated ridges separated by thin-walled integu- 
ment. Ventral segmental papillae lacking or with 2 pairs of large papillae on 
segments 11 and 12, extending dorsally between neuropodia. Pygidium dorsal, 
large, bulbous, wedged between parapodia of posterior 3 segments, with pair of 
ventral anal cirri. Pharynx with 7 pairs of papillae—7 dorsal with median one 
larger and 7 ventral with median one short; 2 pairs of jaws with few (5—7) basal 
teeth. 

Etymology.—The generic name is derived from Greek, /epidos, scale or bract; 
notos, back; podos, foot, referring to the bracts on the notopodia. Gender: mas- 
culine. The specific name is derived from Latin, fimbria, fringe, referring to the 
fimbriated parapodia. 


Lepidonotopodium fimbriatum, new species 
Figs. 1—5 


Material examined.—Pacific Ocean off western Mexico, 20°50’N, 109°06’W, 
2600 m, Alvin dive 1213, sample B21, 19 April 1982, Clam Acres—holotype, 
USNM 80113, and 2 paratypes, USNM 80114, 80115. Same area, Alvin dive 
1223, sample 10A, 7 May 1982—paratype, young, USNM 80116. 

Description.—The 3 adult types are similar in size: 23-24 mm in length, 12-13 
mm in width, including setae, with 28 segments, the last segment very small. All 
specimens are uniformly dark greyish in color, stout, short-bodied, rectangular 
in outline, flattened ventrally, strongly arched dorsally, and slightly tapered and 
rounded anteriorly and posteriorly (Fig. 1). The elytra are thick, leathery, imbri- 


394 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 2. Lepidonotopodium fimbriatum, paratype, USNM 80115: A, Anterior end, dorsal view; B, 
Anterior end, ventral view; C, Left side of segments 12 (cirrigerous) and 13 (elytrigerous), dorsal 
view; D, left side of posterior end (segments 21-27 and pygidium), dorsal view; E, Left first elytron, 
place of attachment dotted; F, Left second elytron; G, Left middle elytron; H, Right eleventh elytron. 
Scales = 1.0 mm for A—D; 1.0 mm for E-H. 


VOLUME 96, NUMBER 3 395 


Fig. 3. Lepidonotopodium fimbriatum, paratypes, A, USNM 80114; B-1, USNM 80115: A, Left 
side of segments 10-13, ventral view; distal tip of segmental papilla extending dorsally between 
parapodia shown separately; B, Distal part of pharynx cut open and spread apart, ventral half on left, 
dorsal half on right; C, Dorsal jaws and median dorsal papilla; D, Tentacular parapodium, inner view, 
aciculum dotted; E, Elytrigerous parapedium from segment 2, anterior view; F, Notoseta from same; 
G, Neuroseta from same; H, Cirrigerous parapodium from segment 3, posterior view; I, Elytrigerous 
parapodium from segment 4, posterior view. Scales = 1.0 mm for_A, B; 0.5 mm for C; 1.0 mm for 
D, E, H, I; 0.1 mm for F, G. 


cated, covering the dorsum (Fig. 1). There are 11 pairs of elytra attached on 
segments 2, 4, 5, 7, 9, 11, 13, 15, 17, 19 and 21, with dorsal cirri on the posterior 
7 segments (Fig. 2D). Except for the first and last pairs, each elytron has 2 raised 
smooth macrotubercles on the posterior one-third, not sharply set off from the 
surface (Figs. 1, 2F, G). The first pair of elytra (on segment 2) have an additional 
tubercle on the medial side of the place of attachment (Fig. 2E). The last pair of 
elytra (on segment 21) lack macrotubercles; they are more elongated and cover 
the posterior cirrigerous segments and pygidium (Fig. 2H). The elytra are reni- 


396 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


———}I 


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Fig. 4. Lepidonotopodium fimbriatum, paratype, USNM 80115: A, Left middle elytrigerous para- 
podium, anterior view; B, Left middle cirrigerous parapodium, posterior view; C, Enlarged notopo- 
dium and bract, posterior view, notoaciculum dotted; D, Enlarged distal part of neuropodium, pos- 
terior view, neuroaciculum dotted; E, Notosetae from middle parapodium; F, Upper, middle and 
lower neurosetae from middle parapodium. Scales = 1.0 mm for A, B; 0.5 mm for C, D; 0.1 mm for 
E, F. 


form, attached eccentrically to large prominent elytrophores (Figs. 2A, C, D, 31, 
4A). The elytral surface appears smooth but is covered with numerous round 
microtubercles giving a dotted appearance, along with some scattered globular 
micropapillae. The dorsal cirri on the segments lacking elytra have cylindrical 
cirrophores attached dorsoposteriorly on the notopodia; they are wider basally 
with a bulbous lobe on the posterior side; the styles are tapering and extend to 
about the tips of the neurosetae (Figs. 2A, C, 3H, 4B). The dorsal tubercles on 


VOLUME 96, NUMBER 3 397 


Fig. 5. Lepidonotopodium fibriatum, young paratype, USNM 80116: A, Anterior end, dorsal view; 
B, Right parapodia of segments 5—7, dorsal view; C, Posterior end, dorsal view; D, Notosetae; E, 
Neurosetae. Scales = 0.2 mm for A-C; 0.1 mm for D, E. 


the cirrigerous segments, corresponding in position to the elytrophores, are large 
and inflated. The surfaces of both the elytrophores and dorsal tubercles, including 
their bases, have numerous ciliated ridges interspersed with thin integument, 
sometimes appearing as pits (Figs. 2C, D, 3H, I, 4A, B). The ridges continue on 
the upper halves of the anterior and posterior sides of the parapodia altogether 
forming well-developed ciliated lacunar areas. The surface of the middorsum is 
wrinkled, that of the ventral surface essentially smooth. 

The prostomium is bilobed, the anterior lobes subtriangular, each with a small 
frontal filament; lateral antennae are absent (Fig. 2A, B). The median antenna is 
inserted in the anterior notch, having a short clyindrical ceratophore and short 
subulate style. The palps are cylindrical, smooth, with filamentous tips, about 
twice the length of the prostomium. Eyes are lacking. The first or tentacular 
segment is not visible dorsally; the tentaculophores are lateral to the prostomium, 
each with a single aciculum but without a distinct acicular lobe and without setae 
(Figs. 2A, B, 3D). The styles of the 2 pairs of dorsal and ventral tentacular cirri 
are similar in length, smooth, tapered, slightly shorter than the palps. A facial 
tubercle is lacking. 

The second or buccal segment bears the first pair of large elytrophores, bira- 
mous parapodia and ventral or buccal cirri attached basally on prominent cirro- 
phores lateral to the mouth; their styles are similar to the tentacular cirri and 
longer than the following ventral cirri (Figs. 2A, B, 3E). The ventral mouth is 
enclosed in upper, lateral, and posterior lips between segments | to 3 (Fig. 2B). 
The pharynx was not extended but was dissected out and slit open (Fig. 3B, C). 
The opening of the large muscular pharynx is encircled by 7 pairs of large bulbous 
papillae: 7 dorsal with the median one larger and 7 ventral with the median one 
smaller. The 2 pairs of dorsal and ventral jaws are fused medially and the basal 
parts are serrated with 5-7 teeth. 


398 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


The biramous parapodia have shorter notopodia located on the anterodorsal 
sides of the longer neuropodia (Figs. 3H, I, 4A, B). The notopodia are conical, 
with projecting acicular lobes and enclosed anterodorsally by well-developed large 
flaring bracts (Figs. 2C, D, 4A—C). The neuropodia are diagonally truncated and 
deeply cleft on the upper part. The presetal acicular lobe projects dorsally beyond 
the rounded postsetal lobe; the upper margin of the latter has 3 rounded papillated 
extensions (Fig. 4B, D). The distal margins of the notopodial acicular lobes, 
notopodial bracts, and neuropodial lobes are fimbriated with slender papillae. The 
notosetae are numerous, forming radiating bundles; they vary in length, are sim- 
ilar in width to the neurosetae and have relatively few (4-7; 7-11 on segment 2) 
widely-spaced spines along one side and blunt rounded tips (Figs. 3F, 4E). The 
neurosetae are numerous, forming fan-shaped bundles. They are all similar, the 
upper ones being slightly more slender and the lower ones shorter (more slender 
on segment 2); they have 2 rows of numerous spines along one side, with slightly 
hooked bare tips (Figs. 3G, 4F). The ventral cirri are short, tapered, smooth and 
attached on the middle of the neuropodia (Fig. 3A). 

Segmental or nephridial papillae are not obvious. On one of the 3 adult types 
(USNM 80114), there are 2 pairs of large bulbous papillae on the ventral side of 
segments I1 and 12, narrowing and projecting dorsally between the parapodia 
(Fig. 3A). The pygidium is visible dorsally as a bulbous lobe wedged between the 
parapodia of the 3 posterior smaller segments (26—29), the last one being quite 
small; there is a pair of long ventral anal cirri, similar to the posterior dorsal cirri 
(Fig. 2D). 

The small paratype (USNM 80116), collected from the same site but on a later 
dive, is | mm long, | mm wide including setae, with 11 segments plus a growing 
zone and pygidium (Fig. 5C). No elytra remain. The prostomium resembles that 
found in the adults except that the palps are bulbous basally and slender more 
distally (Fig. SA). The tentacular and dorsal cirri are also wider basally (Fig. 5A, 
B). The notosetae have more numerous spines (Fig. 5D); the neurosetae are more 
slender and have fewer and larger spines (Fig. SE). 

Remarks.—Based on the structure of the prostomium and tentacular segment, 
Lepidonotopodium could be placed in the subfamily Macellicephalinae, as revised 
by Pettibone (1976), in having the prostomium bilobed with short frontal filaments 
on the anterior lobes, a median antenna with a distinct ceratophore in the anterior 
notch, without lateral antennae, with paired palps and two pairs of tentacular 
cirri lateral to the prostomium. Of the genera in the Macellicephalinae, Lepido- 
notopodium is closest to Bathykurila Pettibone, based on the type of notosetae 
and neurosetae. The shapes of the parapodia differ, however. 

Jaws with serrated basal plates, characteristic of the aphroditacean family 
Acoetidae (=Polyodontidae), are not the usual type in the Polynoidae but they 
are known for some bathyal species, such as Bathyvitiazia pallida (Levenstein) 
and Bruunilla natalensis Hartman in the Macellicephalinae and for Bathyedithia 
berkeleyi (Levenstein) and B. tuberculata Levenstein in the Bathyedithinae (Pet- 
tibone 1976, 1979; Levenstein 1981). 

Lepidonotopodium 1s unique in the Polynoidae in having well-developed bracts 
encircling the notopodia. This is known in some other families of the scaled 
polychaetes or Aphroditacea, such as the Sigalionidae, but not in the Polynoidae. 

The numerous ciliated ridges separated by thin integument on the elytrophores, 
dorsal tubercles and anterior and posterior sides of the parapodia appear to be 


VOLUME 96, NUMBER 3 399 


unique in the Polynoidae. Beneath the surface there appear to be sponge-like 
areas with ciliated lacunar passages, perhaps serving for respiratory exchange, a 
feature worthy of future investigation. 

The presence of fimbriated parapodia, formed of filiform papillae on the distal 
margins of the notopodia, notopodial bracts, and neuropodia, are also at least 
unusual in the Polynoidae. Filiform papillae are found on the distal margins of 
the neuropodia in some members of the Lepidonotinae, such as Hermenia Grube, 
Halosydnopsis Uschakov and Wu, Parahalosydnopsis Pettibone, and Lepido- 
nopsis Pettibone, as indicated by Pettibone (1975, 1977). 

The type of elytra with nodular macrotubercles not sharply set off from the 
surface, as found in Lepidonotopodium fimbriatum, is also unusual in the Poly- 
noidae. Similar structures are found on the elytra of Bathynoe nodulosa Ditlevsen 
(1917:42, pl. 3: fig. 12), collected in the North Atlantic south of Iceland in a depth 
of 1992 meters, and in Bathynoe nodulosa pacifica Uschakov, 1950, from the 
Okhotsk Sea in a depth of 1366 meters (see Uschakov 1955:134, fig. 24C; 1965S: 
115, fig. 24C, as Weberia). Lepidonotopodium and Bathynoe (=Weberia, preoc- 
cupied) differ in a number of other features. 


Acknowledgments 


My thanks go to the OASIS group of Scripps Institution of Oceanography for 
the privilege of working up this interesting material. The manuscript benefited 
from the reviews by my colleagues Dr. Meredith L. Jones and Dr. Kristian Fau- 
chald. Mr. Michael Carpenter kindly photographed the holotype. 


Literature Cited 


Ditlevsen, Hj. 1917. Annelides I.—The Danish Ingolf-Expedition, Copenhagen 4(4): 1-71, 6 pls. 

Levenstein, R. J. 1981. [Some peculiarities in the distribution of the polychaetes (Family Polynoidae) 
in the Canada basin of the Arctic Ocean.]—Transactions of the P. P. Shirshov Institute of 
Oceanology 115:26—36, figs. 1-4. [In Russian, English summary. ] 

Pettibone, M. H. 1975. Review of the genus Hermenia, with a description of a new species (Poly- 
chaeta: Polynoidae: Lepidonotinae).—Proceedings of the Biological Society of Washington 
88(22): 233-248, figs. 1-5. 

—.. 1976. Revision of the genus Macellicephala McIntosh and the subfamily Macellicephalinae 
Hartmann-Schroder (Polychaeta: Polynoidae).—Smithsonian Contributions to Zoology 229: 1— 
71, figs. 1-36. 

——. 1977. Review of Halosydnopsis and related genera (Polychaeta: Polynoidae: Lepidonoti- 
nae). In Essays on polychaetous annelids in memory of Dr. Olga Hartman, D. J. Reish and 
K. Fauchald, editors.—Allan Hancock Foundation, University of Southern California, pp. 39— 
62, figs. 1-6. 

——. 1979. Redescription of Bruunilla natalensis Hartman (Polychaeta: Polynoidae), originally 
referred to Fauveliopsidae.—Proceedings of the Biological Society of Washington 92(2):384— 
388, figs. 1-2. 

Uschakov, P. V. 1950. [Polychaete Worms (Polychaeta) from the Okhotsk Sea.|—Issledovaniia 
Dalnevost Morei SSSR 2:140—234, 39 figs., 2 pls. [In Russian. ] 

——. 1955. [Polychaeta of the Far Eastern Seas of the U.S.S.R.] In Keys to the fauna of the 
U.S.S.R.—Zoological Institute of the Academy of Sciences of the U.S.S.R. No. 56:1—445, 
164 figs. [In Russian; 1965. English translation by Israel Program for Scientific Translations, 
419 pp.] 


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


PROC. BIOL. SOC. WASH. 
96(3), 1983, pp. 400-406 


MINUSCULISQUAMA HUGHESI, A NEW 
GENUS AND SPECIES OF SCALE WORM 
(POLYCHAETA: POLYNOIDAE) FROM 
EASTERN CANADA 


Marian H. Pettibone 


Abstract.—A new polynoid polychaete, Minusculisquama hughesi n. gen., n. 
sp., from Northumberland Strait and Nova Scotia is described. The species is 
remarkable for its elongate and flattened body with only 15 pairs of minute elytra 
confined to the anterior region. It shows features characteristic of some polynoids 
found living commensally in the tubes of other polychaetes. 


Some polynoids from Eastern Canada were sent to me for identification from 
two sources. A single specimen from Northumberland Strait, off Prince Edward 
Island, collected by J. F. Caddy et al. in 1975, was sent to me by Mrs. Leslie E. 
Linkletter of the Biological Station in St. Andrews, New Brunswick. It was col- 
lected at station 95 listed by Caddy et al. (1977:9). Nine specimens were collected 
in St. Margaret’s Bay, Nova Scotia, by T. G. Hughes in 1976 and sent to me by 
Mrs. Patricia Pocklington of the Nova Scotia Museum. They were collected at 
station 1 reported by Hughes (1979:530). The specimens proved to belong to a 
new genus and species. 

The types are deposited in the Biological Station at St. Andrews, New Bruns- 
wick (BSNB), the National Museum of Canada, Ottawa (NMC), the Nova Scotia 
Museum, Halifax (NSM) and the National Museum of Natural History, Smith- 
sonian Institution, Washington, D.C. (USNM). 


Family Polynoidae 
Minusculisquama, new genus 


Type-species.—Minusculisquama hughesi, new species. 

Diagnosis.—Body elongate, with numerous segments (about 100). Elytra 15 
pairs, minute, on slightly developed elytrophores on segments 2, 4, 5, 7, alternate 
segments to 23, 26, 29, and 32. Dorsal cirri on non-elytra-bearing segments. Dor- 
sal tubercles indistinct. Prostomium bilobed, with small cephalic peaks, 3 anten- 
nae, paired palps and 2 pairs of eyes. Median antenna with distinct ceratophore 
in anterior notch, lateral antennae with ceratophores inserted ventrally and con- 
verging midventrally. Tentaculophores of first or tentacular segment lateral to 
prostomium, achaetous, with 2 pairs of tentacular cirri; without facial tubercle. 
Second or buccal segment with first pair of elytra, setigerous parapodia and ven- 
tral buccal cirri; without nuchal fold. Parapodia subbiramous, notopodia small, 
with notoaciculum only, without notosetae (rarely single one present). Neuro- 
podia with rounded anterior acicular and posterior lobes, deeply cut dorsally and 
ventrally, without projecting acicular lobes. Neurosetae stout, relatively few, with 
few spines, tapering to slender tips; upper ones stouter, smooth, with blunt tips. 


VOLUME 96, NUMBER 3 401 


Pygidium with pair of anal cirri. Nephridial papillae small, beginning on segment 
6. Pharynx with 9 pairs of papillae and 2 pairs of jaws. 

Etymology.—The generic name is derived from the Latin, minusculus, small, 
squama, scale, referring to the small scales or elytra. Gender: feminine. The 
species is named for Dr. Trevor G. Hughes, one of the collectors. 


Minusculisquama hughesi, new species 
Figs. 1-3 


Material examined.—EASTERN CANADA: Northumberland Strait off Prince 
Edward Island, 46°19'20’N, 62°10'10"W, 34 m, red mud over clay, sta. 95, 23 July 
1975, J. F. Caddy et al., collectors—paratype, BSNB 2290. St. Margaret’s Bay, 
Nova Scotia, 44°32'’N, 63°59’W, 70 m, silty mud and sand, sta. 1, 28 August 1976, 
T. G. Hughes, collector—holotype, USNM 72887, 3 paratypes, USNM 72888, 2 
paratypes, NSM 976Z351.49 and 3 paratypes, NMC 1982-0885-7. 

Measurements.—The holotype from St. Margaret’s Bay (USNM 72887) is 98 
mm long, 7.5 mm wide, including setae, with 108 segments. The length of 3 
paratypes from the same collection (USNM 72888) are 66-89 mm, widths 7-8 
mm, with 76-113 segments. The single paratype from Northumberland Strait 
(BSNB 2290) has a length of 46 mm, a width of 5 mm, including setae, with 58 
segments plus a small regenerating posterior end of about 10 segments. 

Description.—The body is elongate, flattened, of about equal width along the 
body, tapering slightly anteriorly and posteriorly, with a rather deep midventral 
longitudinal groove. The body is dusky dorsally, especially on the posterior half 
and ventrally along the rather deep midventral longitudinal groove and extending 
laterally as transverse bands. 

There are 15 pairs of minute nipple-like elytra on segments 2, 4, 5, 7, alternate 
segments to 23, 26, 29 and 32. They are not the usual polynoid type and are easily 
overlooked (Figs. 1A, C, D, 2B). They are somewhat flattened, disc-like, attached 
to rather indistinct, slightly bulbous elytrophores. They lack tubercles; some 
micropapillae are present on the surface (Fig. 2B). Dorsal cirri are present on the 
rest of the segments (Figs. 1A, C, F, 2A, D, E, 3A, B, D). Dorsal tubercles on 
the cirrigerous segments are indistinct. 

The prostomium is bilobed with small cephalic peaks (Fig. !A—C). The median 
antenna has a large ceratophore in the anterior notch, with a style of about the 
same length as the prostomium, with a tapered tip. The lateral antennae have 
small ceratophores inserted ventrally and converge midventrally (Fig. 1B); the 
styles are subulate and about a third as long as the median antenna. The palps 
are stout, tapered, slightly longer than the median antenna. The 2 pairs of eyes 
are rather small, the anterior pair in the region of greatest prostomial width is 
slightly larger than the posterior pair. The tentaculophores of the achaetous first 
or tentacular segment are lateral to the prostomium, with 2 pairs of tentacular 
cirri similar to the median antenna; the dorsal tentacular cirri are about equal in 
length to the median antenna, the ventral ones slightly shorter. There is no facial 
tubercle. Segment 2 bears the first pair of small elytra, subbiramous parapodia 
and ventral buccal cirri that are slightly longer than the following ventral cirri 
(Fig. 1A—D). Notosetae are lacking. The neurosetae are similar to the following 


402 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 1. Minusculisquama hughesi, A, B, paratype, BSNB 2290; C—G, holotype, USNM 72887: 
A, Anterior end, dorsal view, styles of left dorsal tentacular cirrus and right dorsal cirrus from segment 
3 missing; B, Anterior end, ventral view, pharynx partially extended; C, Anterior end, dorsal view, 
left dorsal tentacular cirrus shorter, regenerating; D, Right elytrigerous parapodium from segment 2, 
anterior view, acicula dotted; E, Upper, middle and lower neurosetae from same; F, Right cirrigerous 
parapodium from segment 3, posterior view; G, Upper, middle and lower neurosetae from same. 
Scales: = 1.0 mm for A—C; 0.5 mm for D, F; 0.1 mm for E, G. 


ones except that they are more slender (Fig. 1E). The ventral mouth is enclosed 
in the anterior 3 segments (Fig. 1B). 

The parapodia are subbiramous and similar along the length of the long body 
(Figs. 1[A—D, F, 2A, B, D, E, 3A, B, D). The notopodia are small, subconical, 
located on the anterodorsal bases of the neuropodia, each with a notoaciculum 
only. Notosetae are lacking. However, a single notoseta was found on segment 


VOLUME 96, NUMBER 3 403 


Fig. 2. Minusculisquama hughesi, holotype, USNM 72887: A, Right cirrigerous parapodium from 
segment 12, posterior view, micropapillae shown separately; B, Right elytrigerous parapodium from 
segment 13, anterior view, elytral micropapillae shown separately, acicula dotted; C, Upper, middle 
and lower neurosetae from same; D, Right middle parapodium (about segment 64), posterior view; 
E, Same, anterior view; F, Upper, middle and lower neurosetae from same. Scales: = 0.5 mm for 
A, B, D, E; 0.1 mm for C, F. 


404 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 3. Minusculisquama hughesi, A-C, holotype, USNM 72887; D-F, paratype, USNM 72888; A, 
Right parapodium from posterior region (about segment 84), posterior view; B, Same, anterior view; 
C, Upper, middle and lower neurosetae from same; D, Right parapodium from segment 85, posterior 
view; E, Notoseta and distal tip of notopodium from same; F, Upper, middle and lower neurosetae 
from same. Scales: = 0.5 mm for A, B, D; 0.1 mm for C, E, F. 


85 on one of the paratypes (USNM 72888). It is slender, with 6 spinous rows, and 
tapers to a slender tip (Fig. 3D, E). The neuropodia are elongate, deeply cut 
dorsally and ventrally. The presetal acicular lobes are larger, diagonally truncate 
on the lower half, truncate on the upper half; the postsetal lobes are smaller, 
subconical and rounded, the upper part straight forming a deep notch with the 
anterior lobe. The neurosetae are rather stout and relatively few in number (6— 
13). They are wider subdistally and taper to pointed tips, not hooked (Figs. 1E, 
G, 2C, F, 3C, F). They are variable in size, the upper ones (1—3) are stouter, 
darker and smooth, with blunt tips (perhaps worn). The middle and lower neu- 
rosetae have a few spines on the wider part (2-4 pairs). The spines may be broken 
off with only their bases evident. The lower posterior group of neurosetae (4—5) 


VOLUME 96, NUMBER 3 405 


are shorter and more slender. The dorsal cirri have cylindrical cirrophores at- 
tached posterodorsally on the notopodia; the styles are short, tapering and extend 
to about the tips of the neuropodia or beyond. The styles have scattered micro- 
papillae (Fig. 2A). The ventral cirri, attached on the middle of the neuropodia, 
are short, thick and subulate, smooth or with scattered micropapillae. 

The muscular pharynx has the usual 9 pairs of papillae and 2 pairs of jaws. 
The nephridial papillae begin on segment 6; they are short, cylindrical, extending 
posterolaterally. The pygidium is small, rounded, with a pair of anal cirri, longer 
than the dorsal cirri. 

Biology.—Minusculisquama hughesi was collected in silty mud and sand in 70 
meters in St. Margaret’s Bay and in red clay over mud in 34 meters in North- 
umberland Strait. Although there is no data with the specimens to confirm it, 
their structure suggests that they were probably living commensally in the tubes 
of other polychaetes, such as maldanids. In his report on the collections from St. 
Margaret's Bay, Hughes (1979:531) indicated that “‘the top 20 cm of sediment 
were permeated by numerous tubes made by various species of maldanid poly- 
chaetes. Some of the tubes protruded above the surface of the sediment.’ I 
suggest that the host of M. hughesi may very well have been a maldanid. Com- 
mensalism is suggested by the following features of M. hughesi: the elongate 
flattened body of about equal width, the small prostomial eyes, the minute elytra 
confined to the anterior region, the reduced notopodia and missing notosetae, the 
enlarged upper neurosetae, such as are found in Lepidametria commensalis. 

Minusculisquama agrees with Arctonoe Chamberlin, 1920, and Adyte Saint- 
Joseph, 1899, in that all three have elongated bodies with numerous segments, 
similar type of prostomia with the lateral antennae having distinct ceratophores 
inserted terminoventrally, and subbiramous parapodia with the neuropodia deeply 
cut dorsally and ventrally forming rounded presetal acicular and postsetal lobes. 
In Arctonoe, the elytra are large, more numerous, continuing to the posterior end 
of the long body, and with a different arrangement; also the neurosetae are strong- 
ly hooked, rather than tapered (see Pettibone 1953:56—66). Minusculisquama agrees 
further with Adyte in having 15 pairs of elytra confined to the anterior region of 
the long body and in similar arrangement. In Adyfe, notosetae are present rather 
than absent, the elytra are large and attached to prominent elytrophores rather 
than minute with indistinct elytrophores, and the neurosetae have basal semilunar 
pockets and the upper neurosetae are not larger, as they are in Minusculisquama 
(see Pettibone 1969:5-8). 

Minusculisquama hughesi resembles Lepidasthenia accolus Estcourt, 1967, 
commensal in the burrow of an arenicolid polychaete, in having very small elytra, 
an elongated body, similar prostomia and neuropodia and in lacking notosetae. 
In L. accolus, the elytra are more numerous and arranged on segments 2, 4, 5, 7, 
continuing on alternate segments to the end of the long body; the presetal acicular 
lobes of the neuropodia are notched distally and the dorsum is finely papillated 
(see Estcourt 1967:68—69). 


Acknowledgments 


My thanks go to Leslie E. Linkletter of the Biological Station in St. Andrews 
and Patricia Pocklington of the Nova Scotia Museum for sending me the polynoid 


406 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


specimens, as well as furnishing data regarding their collection. My colleagues, 
Meredith L. Jones and Kristian Fauchald, kindly reviewed the manuscript. 


Literature Cited 


Caddy, J. F., T. Amaratunga, M. J. Dadswell, T. Edelstein, L. E. Linkletter, B. R. McMullin, A. 
B. Stasko, and H. W. vande Poll. 1977. 1975 Northumberland Strait Project, Part I: Benthic 
fauna, flora, demersal fish and sedimentary data.—Fisheries and Marine Service Manuscript 
Report 1431, The Biological Station, St. Andrews, New Brunswick, pp. 1-46. 

Estcourt, I. N. 1967. Burrowing polychaete worms from a New Zealand estuary.—Transactions of 
the Royal Society of New Zealand 9:65-78, 10 figs. 

Hughes, T. G. 1979. Studies on the sediment of St. Margaret's Bay, Nova Scotia.—Journal of the 
Fisheries Research Board of Canada 36:529-536. 

Pettibone, M. H. 1953. Some scale-bearing polychaetes of Puget Sound and adjacent waters.— 
University of Washington Press, Seattle, 89 pp., 40 pls. 

——. 1969. Review of some species referred to Scalisetosus McIntosh (Polychaeta, Polynoi- 
dae).—Proceedings of the Biological Society of Washington 82:1—30, 12 figs. 


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


PROC. BIOL. SOC. WASH. 
96(3), 1983, pp. 407-410 


ARCTODRILUS WULIKENSIS, NEW GENUS, 
NEW SPECIES (OLIGOCHAETA: TUBIFICIDAE) 
FROM ALASKA 


Ralph O. Brinkhurst and R. Deedee Kathman 


Abstract.—Arctodrilus is a new tubificine tubificid genus with hair and pecti- 
nate setae but no genital setae, male ducts with vasa deferentia about as long as 
the atria and penes plus ejaculatory ducts, and atria spindle-shaped with median 
prostates. The type (and only) species A. wulikensis from the Wulik River system 
in Alaska has all the postclitellar setae with long upper teeth. In the bifid anterior 
ventrals and in the pectinate dorsals the teeth are more nearly equal; the hair and 
pectinate setae occur from II to XIV. Psammoryctides hadzii Karaman has male 
ducts like those of Arctodrilus (which are very general in form) but it does have 
spermathecal setae. While it is not (as described) a typical member of Psam- 
moryctides, P. hadzii is left incertae sedis in that genus pending a revision of the 
species based on examination of the type-material. 


During an environmental investigation into natural mineral seepages into the 
Wulik River system headwaters, a new tubificine tubificid was discovered that 
cannot be attributed to any known genus. The species was only found at locations 
not affected by the seepages. 


Arctodrilus, new genus 


Definition.—TVubificine tubificid with male ducts having vas deferens little long- 
er than atrium and ejaculatory duct combined, entering atrium apically, atrium 
spindle-shaped with median prostate, ejaculatory duct 1.5 times the length of the 
atrium, of one width and histological composition throughout. Penis present, with 
thin cuticular sheath no thicker than cuticularized penis sac wall. No modified 
genital setae. Hair and pectinate setae present in at least some species. 

Type-species.—Arctodrilus wulikensis, new species, by monotypy. 

Etymology.—Found in Alaska north of the Arctic circle. 

Discussion.—The male ducts of the single new species are simple unspecialized 
structures. The most similar male ducts are those of Limnodrilus Claparéde and 
Spirosperma Eisen species (sensu Brinkhurst 1981) but there are no other simi- 
larities, and the body wall, setae, prostomium, spermathecae and penes differ 
from those of both genera. The genus Tubifex Lamarck lacks ejaculatory ducts, 
and while some species currently included within the genus have the vas deferens 
entering the atrium apically and unassociated with the prostate stalk (see T. 
superiorensis (Brinkhurst and Cook) sensu Brinkhurst 1981), the type and other 
characteristic Tubifex sensu strictu species have the vas deferens and prostate 
Stalk close together under the curved head of a broad, comma-shaped atrium. All 
species in the genus Isochaetides Hrabé emm. Brinkhurst 1981, have very long 
vasa deferentia, and lack hair and pectinate setae, but most have modified genital 
setae. 


408 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


While it seems surprising at first glance that an animal with such a simple male 
duct cannot be attributed to an existing genus, this new species cannot even be 
attributed to the genus Twbifex sensu latu, and to do so would simply defer the 
decision that is taken here. 

Distribution.—Alaska, Wulik River watershed, DeLong Mountains above Kiv- 
alina (see species description). 


Arctodrilus wulikensis, new species 
Fig. 1 


Description.—Up to 43 segments, length 8.0 mm, width 0.2 mm. Dorsal setae 
anteriorly 2-4 or even 5 long serrate hairs, with serrations probably on one side; 
thin and as long as body width; 2-4 or 5 pectinate setae which, under lower power 
magnification, appear to have long, thin outer teeth and thin intermediate teeth 
arranged in a broad, semi-palmate form but which appear more typically pectinate 
under oil immersion lens (Fig. 1). Hair setae and pectinate setae absent beyond 
XIV, where they are replaced by 2-3 bifid setae resembling the ventral setae. 
Ventral anterior setae 3—S per bundle, bifid with upper teeth thinner than but only 
a little longer than the lower, behind XIV with upper tooth thinner but twice as 
long as the lower; no modified genital setae. Reproductive system as for the 
Tubificinae, with all organs paired. Spermatheca with short stout duct, volumi- 
nous and thin-walled ampulla and lateral pore. Spermatozeugmata present, broad. 
Male duct with thin vas deferens which widens distally but narrows immediately 
prior to insertion at apex of atrium; atrium tubular, widest just above the mid- 
point where a substantial stalked prostate enters. A substantial ejaculatory duct 
connects the atrium with the penis sac; the penis sac is ovoid-oblong with cutic- 
ular walls, the contained penis is of similar shape and has a thin cuticular sheath 
no thicker than the cuticle on the penis sac wall. The penis sac may be eversible, 
but there is a true penis within the sac (Fig. 1). 

Type-locality.—Red Dog Creek and Ikalukrok Creek, part of Wulik River sys- 
tem, DeLong Mountains, Brooks Range, Alaska (68°05'N, 162°45’W), in areas 
not affected by seepages containing large amounts of zinc, cadmium, iron, or 
manganese. July-August 1982. 

Etymology.—From the Wulik River drainage. 

Holotype.—USNM 80444, a dissected whole mount in Canada Balsam. 

Paratypes.—USNM 80445-80453, 7 whole mounts and 70 specimens in fluid. 
Brinkhurst collection 5 whole mounts, Kathman collection 2 whole mounts. 

Discussion.—As with the generic diagnosis, there is little of a specialized nature 
to comment upon in the species description. The setation is somewhat unusual 
in that the upper teeth of the dorsal and ventral postclitellar setae are longer than 
the lower teeth, whereas the opposite trend is more usual. A similar tendency to 
elongation of the upper tooth posteriad is seen in the naidid genus Amphichaeta 
Tauber, but in the new taxon considered here the change from one form to the 
other is abrupt and coincides with the loss of hair and pectinate setae dorsally. 
The tendency to loss of hair and pectinate setae posteriad is common in tubificids 
but is usually progressive, with replacement of pectinates by bifids of essentially 
similar shape occurring before shortening and reduction in number or total loss 
of hair setae. 


VOLUME 96, NUMBER 3 409 


MALE DUCT 
A. WULIKENSIS 


SPERMATHECA 


Vas Deferens 


Atrium 


Prostate 


Ejaculatory Duct 


Hair Pectinate V 


Pectinate if ih 
MALE DUCT MALE DUCT 


Anterior 
P. HADZII P. MORAVICUS Ventral Ventral XV 


Fig. 1. A. wulikensis, reproductive structures and setae, from holotype and paratype respectively; 
Psammoryctides hadzii and P. moravicus, male ducts, redrawn from literature in comparable position. 


We are aware of only one other species with male ducts that are supposed to 
resemble those of A. wulikensis to some extent, and that is Psammoryctides 
hadzii, described by Karaman (1974). According to the illustration (Fig. 1), that 
species cannot be attributed to Psammoryctides as there is no enlargement of the 
ejaculatory duct and the atrium is not the usual small globular body, both essential 
characteristics of that genus. Psammoryctides hadzii is discussed by its author 
in relation to P. ochridanus (Hrabé), although that species has male ducts and 
spermathecal setae of the form typical of the genus and P. hadzii has only the 
latter. Similar spermathecal setae are found in other genera, however, so this 
character by itself is not diagnostic. There remains the very unlikely possibility 


410 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


that the male ducts of Psammoryctides species resemble those of Arctodrilus 
prior to full maturation or during the early stages of post copulatory resorption. 
Specimens of A. wulikensis had sperm in the spermathecae, and while the male 
ducts are of a delicate construction, they do appear to be fully formed. No men- 
tion of spermatozeugmata appears in the description of P. hadzii and so the degree 
of maturation of these worms from an underground river in Jugoslavia is un- 
known. We conclude that no useful purpose would be served by placing P. hadzii 
in Arctodrilus until the latter is better known, especially as the former does 
possess genital setae, and the zoogeographic evidence supports that decision. 
Psammoryctides hadzii should remain a species incertae sedis of Psammoryc- 
tides. 

Five specimens of A. wulikensis were collected in early July, 12 were found in 
late July, and 68 were obtained in late August, in both Red Dog and Ikalukrok 
creeks. The lumbriculid Rhynchelmis brooksi Holmquist and unidentified En- 
chytraeidae were also collected with our specimens. All were found in cold, fast 
riffle areas characterized by cobble/pebble/gravel substrate. Other dominant mac- 
rofauna included Ephemeroptera, Plecoptera, Simuliidae, and Chironomidae. 


Acknowledgments 


The authors are indebted to E.V.S. Consultants Limited and to the Alaska 
Department of Environmental Conservation for access to this material and per- 
mission to publish a description of it. The illustration was prepared by Ian 
McSorley. 


Literature Cited 


Brinkhurst, R. O. 1981. A contribution to the taxonomy of the Tubificinae.—Proceedings of the 
Biological Society of Washington 94: 1048-1067. 
Karaman, S. 1974. Beitrag zur Kenntnis der Stisswasseroligochaeten Sloweniens.—Bioloski vestnik 


Institute of Ocean Sciences, P. O. Box 6000, 9860 West Saanich Road, Sidney, 
British Columbia V8L 4B2, Canada. 


PROC. BIOL. SOC. WASH. 
96(3), 1983, pp. 411-419 


A CONTRIBUTION TO THE TAXONOMY OF THE 
ENCHYTRAEIDAE (OLIGOCHAETA). REVIEW 
OF STEPHENSONIELLA, WITH NEW 
SPECIES RECORDS 


Kathryn A. Coates 


Abstract.—The marine enchytraeid genus Stephensoniella Cernosvitov, 1934, 
(Oligochaeta: Annelida) is revised. Type-materials of four nominate species, S. 
marina (Moore, 1902), S. barkudensis (Stephenson, 1915), Lumbricillus sterreri 
Lasserre and Erséus, 1976, and Marionina trevori Coates, 1980, were examined. 
Stephensoniella barkudensis possesses a spermatheca with the same diverticulate 
ampullar structure as, and is a junior synonym of, S. marina, the type-species of 
the genus. Lumbricillus sterreri and Marionina trevori are transferred to Ste- 
phensoniella because they possess large, paired, unlobed seminal vesicles; ne- 
phridia with only the funnels in front of the septa; diverticulate spermathecal 
ampullae; thickened septa anteriorly; and only two or three setae in each setal 
bundle. Stephensoniella is structurally close to Marionina and Lumbricillus ; how- 
ever, a conservative evaluation of nephridial, spermathecal, and seminal vesicular 
characteristics distinguishes the species of each. 


Stephensoniella was erected by Cernosvitov (1934) based on a redescription of 
the type-species, Enchytraeus marinus Moore, 1902. New material was collected 
from the type-locality, Gilson Hil Tip, Coney Sol, Bermuda, at the time of the 
redescription because the original material had been lost. Cernosvitov (1934) also 
transferred Enchytraeus barkudensis Stephenson, 1915 to Stephensoniella and 
these have remained the only recognized species. The major generic characters 
used by Cernosvitov (1934) to separate Stephensoniella from the closely related 
Lumbricillus and Marionina were the unlobed form of the seminal vesicles, the 
shape and distribution of the setae, and the diverticulate form of the spermathecal 
ampulla; and from Enchytraeus, the absence of peptonephridia and compact form 
of the penial bulb. 

Nielsen and Christensen (1959) indicated that both species of Stephensoniella 
could be referred to Marionina but their unfamiliarity with the species caused 
them to refrain. They did point out, however, that Marionina is very heteroge- 
neous, including several groups of species with only remote relations among 
them. Lasserre and Erséus (1976) echoed the view of Cernosvitov regarding the 
affinities of Stephensoniella and, although they could not clarify the genus defi- 
nition, preferred to conserve it intact. 

The examination of specimens of S. marina collected and identified by C. 
Erséus and the comparison of these with specimens of Marionina trevori Coates, 
1980, revealed numerous structural similarities. Structural characteristics of Lum- 
bricillus sterreri Lasserre and Erséus, 1976, brought to my attention by C. Erséus, 
led to the reexamination of that species and to the subsequent transferral of both 
these species to Stephensoniella. 


412 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


I have attempted in the following systematic section to establish a homogeneous 
species grouping and to clarify those characteristics that distinguish Stephenso- 
niella. All the described species that could be determined undoubtedly to belong 
in Stephensoniella have been included. 


Materials 


Type and additional material of the four species Stephensoniella marina, S. 
barkudensis, Marionina trevori, and Lumbricillus sterreri were borrowed from 
the British Museum (Natural History) (BMNH); the National Museum of Natural 
Sciences, Canada (NMCIC); the Muséum National d’Histoire Naturelle, Paris, 
France (AH); Smithsonian Institution, National Museum of Natural History 
(USNM); the personal collection of C. Erséus, Sweden (CE); and from EVS 
Consultants Ltd., Sidney, British Columbia (EVS). New, unmounted material 
was Stained in alcoholic borax carmine, dehydrated through an ethanol to xylene 
series and mounted whole in Canada balsam. 


Stephensoniella Cernosvitov, 1934 (emended) 


Type-species.—Enchytraeus marinus Moore, 1902. 

Definition.—Brackish and littoral marine enchytraeids. Setae in 4 bundles from 
II; 2 or 3 setae per bundle in preclitellar segments, 2 (infrequently 3) in intra- and 
postclitellar segments; single-pointed, straight or slightly sigmoid, bent proxi- 
mally. Head pore just anterior to 0/1, other dorsal, coelomic pores absent. 
Oesophageal-intestinal transition gradual, intra- or postclitellar. Peptonephridia, 
oesophageal and intestinal diverticula lacking, but with solid, paired, dorsal, post- 
pharyngeal bulbs. Pharyngeal glands well-developed, broadly united at 3/4, 4/5, 
and 5/6. Dorsal blood vessel origin intra- or postclitellar; dorsal vessel bifurcating 
anterior to brain, in prostomium. Lymphocytes nucleate. Chloragocytes present 
from preclitellar segments. Preseptal part of nephridia includes funnel only; in- 
terstitial tissue around postseptal nephridial canal well-developed; ectal duct ter- 
minal, ventral. Muscle layers of some of septa 6/7 to 9/10 thickened. Ovaries, 
testes and associated ducts and pores all paired, in typical familial positions. 
Seminal vesicles paired, unlobed. Sperm funnels (glandular vasa deferentia) well- 
developed; non-glandular vasa deferentia contained in XII, surrounded medially 
at ectal pores by compact penial bulbs, accessory prostatic glands absent. No 
ventral copulatory glands. Spermathecae paired in V, always attached to, if not 
actually communicating with, oesophagus via short ental ducts. Each ampulla 
with a rounded or ovoid diverticulum. Spermathecal ectal duct with or without 
glands. 

Habitat and distribution.—Coastal marine, usually tropical or subtropical. Ca- 
ribbean Sea; Atlantic Ocean—Bermuda, Florida, and Brazil; Indian Ocean—Bar- 
kuda Island, India; Persian Gulf—Saudi Arabia; northeast Pacific Ocean—British 
Columbia. 

Remarks.—The primary distinguishing characteristics of the genus are the dis- 
tribution of almost straight setae in bundles of only two or three; the thickened, 
muscular, preclitellar septa and well developed pharyngeal glands; the paired, 
unlobed seminal vesicles; and the single, rounded diverticulum found on each 
spermathecal ampulla. Rudimentary peptonephridia (see Stephenson 1915a), also 


VOLUME 96, NUMBER 3 413 


referred to in the literature as solid postpharyngeal bulbs and dorsolateral pha- 
ryngeal appendages, have been reported in species of Marionina (Kossmagk- 
Stephan 1983) and Fridericia, Mesenchytraeus, Lumbricillus, Bryodrilus, Buch- 
holzia, and Henlea (Stephenson 1930) and are therefore not a good distinguishing 
characteristic. 

Cernosvitov (1934) discussed the significance to the generic definition of 
Stephensoniella of the two-layered longitudinal epidermal musculature that he 
observed in §. marina. Stephenson (1930) had previously remarked that a two- 
layered longitudinal musculature occurred frequently in Enchytraeus and Fride- 
ricia. Later, Cernosvitov (1937) pointed out that two types of longitudinal muscle 
fibres (round and ribbon-like) were found in some species of Achaeta, Guarani- 
drilus, Hemienchytraeus, and Fridericia. These genera were not considered to 
be phylogenetically close and were placed (Cernosvitov 1937) in three different 
subfamilies. With only a limited amount of information about this musculature 
character and knowledge that the same arrangement occurs in phylogenetically 
dissimilar groups it is not possible to ascertain the taxonomic significance of the 
characteristic. 

The species of Stephensoniella can be distinguished from each other by their 
different absolute sizes and by the distribution of gland cells around the sper- 
mathecal ectal ducts. 


Stephensoniella marina (Moore, 1902) (emended) 
aN. Figs. 1-3 


Enchytraeus marinus Moore, 1902:80-82. 

Stephensoniella marina (Moore).—Cernosvitov, 1934:233—237, 242-243; Cer- 
nosvitov, 1935:8. 

Stephensionella marina (Moore).—Lasserre and Erséus, 1976:455. 

Enchytraeus barkudensis Stephenson, 1915b:142.—Stephenson 1915a:45-47; 1923: 
113-114. 

Stephensoniella barkudensis (Stephenson)—Cernosvitov, 1934:242—243. 

Stephensoniella barcudensis (Stephenson)—Cernosvitov, 1937:291. 


Material examined.—BMNH 1949.3.1.1010, whole mount (originally labelled 
by Cernosvitov as Enchytraeus neotropicus n. sp. [a name which was never 
subsequently published], Mus. Paris #3764), BMNH 1949.3.1.1011, 3 slides of a 
transversely sectioned specimen, one slide of a longitudinally sectioned specimen, 
French Guyana (see Cernosvitov 1935); BMNH 1949.3.1.1012-1014, 4 whole 
mounts, 3 slides, immature, BMNH 1949.3.1.1015, 3 slides of a longitudinally 
sectioned specimen, collected by J. A. G. Wheeler, Gilson Hil Tip, Coney Sol, 
Bermudas, 8 Feb 1933 (see Cernosvitov 1934); CE 22H, 2 whole mounts, partially 
mature, collected by P. Lasserre, Coney Island, Bermuda, 1973 (see Lasserre 
and Erséus 1976); CE M/77-1, one slide of a longitudinally sectioned specimen, 
CE M77-2, one whole mount and 2 mounted, partly dissected specimens, 2 slides, 
collected by C. Erséus, Virginia Key, Florida, 15 Nov 1977; and EVS, 17 whole 
mounted specimens, collected by Saudi Arabian Tetra Tech Inc., Persian Gulf, 
Saudi Arabia, 1981-1982. 

As §. barkudensis: BMNH 1933.5.25.348, one slide of a longitudinally sec- 
tioned specimen, and BMNH 1933.5.25.351, one slide of a lontidudinally sec- 


414 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 1. Stephensoniella marina, section through anterior segments showing posteriorly distended, 
thickened septa 6/7, 7/8 and 8/9 (from Enchytraeus barkudensis, BMNH 1933.5.25.348). 


tioned specimen, syntypes, Chilka Lake, Barkuda Island, India, collected 1914 
(see Stephenson 1915b); BMNH 1933.5.25.355—357, 3 slides, 2 longitudinally sec- 
tioned specimens, Ennur, Madras, India (see Stephenson 1915a). 

Description.—Length 6 to 10 mm, 46 to 73 segments; diameter to 0.5 mm. 
Cutaneous glands in 3 or 4 transverse rows per segment. Clitellum over XII and 
XIII, glands in numerous, regular transverse rows. Setae straight, 33, 50 to 120, 
usually 80 to 90 wm long; setae of II markedly short, 50 to 60 um; lateral setae 
shorter than ventrals of same segment. Two or 3 setae in anterolateral bundles, 3 
setae in ventral bundles to X or XI, 2 (occasionally 3) setae in postclitellar seg- 
ments. Septa (rarely 6/7) 7/8 to 9/10 thickened (Fig. 1). Lymphocytes abundant 
anteriorly, 20 to 30 wm long. Dorsal blood vessel origin postclitellar, to X XIII. 
Postclitellar segments with intra-segmental muscular bands extending from the 
circular epidermal muscles to muscles of gut. Seminal vesicles extending to 9/10; 
eggs confined to clitellar segments. Sperm funnels 2 to 3 times as long as wide, 
length just less than body width; glands differentially developed around ciliated 
duct; collar 2 or more times as wide as glandular part. Vas deferens 14 to 24 um 
wide, non-muscular. Penial bulbs (Fig. 2A) 53 to 68 wm high, medial to vasa 
deferentia. Male pore simple. Spermathecal pores at 4/5, just ventral to level of 
lateral setae. Ectal ducts of spermathecae 14 to 26 wm wide, 3 to 4 times as long 
as wide; aglandular; communicating basally with ampulla at constriction between 
ampulla and thinner walled, dorsally directed diverticulum. Ampulla approxi- 
mately 50 um wide x 80 um long, diverticulum approximately 30 wm wide x 80 
wm long, more or less ovoid (Fig. 2B—D). Ampullae connecting laterally to oe- 
sophagus in posterior of V. Sperm densely bundled in diverticula, in random 
masses in ampullae. 

Habitat.—Estuarine salt lakes and littoral and supralittoral marine, usually in 
medium coarse sands. With Pontodrilus bermudensis, in India (Stephenson 1915b). 

Distribution.—Barkuda Island, India; French Guyana; Florida, U.S.A.; Ber- 
muda; Persian Gulf, Saudi Arabia. 


VOLUME 96, NUMBER 3 415 


Remarks.—Examination of syntypes of S. barkudensis, BMNH 1933.5.25.348 
and .351, and other specimens from Ennur, Madras, and Cernosvitov’s material 
of S. marina revealed a few misinterpretations in, and omissions from, the liter- 
ature. It was found that the spermathecal ampullae of S. barkudensis (Fig. 2D) 
each bore a basal, ovoid diverticulum as in §. marina (Fig. 2B, C). It is recorded 
at the British Museum (personal communication, E. G. Easton) that their material 
of S. barkudensis originated from the Indian Museum; it was simply labelled 
‘“‘types’’ when received and it is doubtful that a designated holotype remains in 
the Indian Museum. 

Other than a broad geographical separation, partially bridged by the new rec- 
ords from the Persian Gulf, reasons supporting a classificatory distinction of S. 
barkudensis from §. marina are not found (Fig. 3). By seniority, S. marina (Moore, 
1902) has precedence for the specific name; by original designation (Cernosvitov 
1934), it is the type-species of the genus. 


Stephensoniella sterreri (Lasserre and Erséus, 1976), new combination 
Fig. 3 


Lumbricillus sterreri Lasserre and Erséus, 1976:453—454 (partim). 


Material examined.—Holotype: AH 193, a whole mount, Ireland Island, Ber- 
muda, collected by P. Lasserre, 1973. Paratypes: AH 195, 196, 2 whole mounts, 
Coot Pond, Bermuda, collected by P. Lasserre, 1973. 

Description.—Holotype with 47 segments, paratypes with 41 and 39 segments. 
Setae 3 per bundle in at least some preclitellar ventral bundles, 2 in other bundles, 
slightly sigmoid, 50 to 60 um long. Clitellum incomplete ventromedially, between 
male pores. Some of septa 7/8 to 9/10 thickened. Vasa deferentia approximately 
13 wm wide, opening ectally via simple pore, lateral to small penial bulb. Seminal 
vesicles and testes unlobed. Spermathecal ectal duct relatively short, 33 to 43 
pm long, bulbous near middle, 17 to 26 wm wide. Glandular cells originating 
along length of duct, rosette of larger glands originating around ectal pore (Fig. 
3). Spermathecal ampulla ovoid, approximately 51 to 61 wm wide and 86 to 91 
pm long, approximately 1.5 times as long as wide. Subapical diverticulum nearly 
globular, 51 to 54 wm in diameter. 

Remarks.—Characteristics not given in the description are as originally de- 
scribed (Lasserre and Erséus 1976) or as for the genus. The primary character- 
istics distinguishing S. sterreri from the type species (Fig. 3) are its small size 
and its spermathecal structure including the position of the origin of the divertic- 
ulum, relative shapes and sizes of the ampulla and diverticulum, and the presence 
of glandular cells on the ectal duct. Three setae may be present in very few 
anteroventral setal bundles, as in paratype AH 196 with three setae only in X 
and XI. 

The Muséum National d’Histoire Naturelle in Paris had no deposition records 
for paratype AH 197 (J. Renaud-Mornant, personal communication) (see Lasserre 
and Erséus 1976). Paratype AH 194 has been referred to S. trevori (see page 
416). 

Habitat.—Mid-littoral in medium or coarse sands rich with organic debris. 

Distribution.—Bermuda. 


416 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


50 um 


Fig. 2. Stephensoniella marina. A, Longitudinal section of glandular penial bulb (p) in XII, vas 
deferens (Vv) opens into a narrow epidermal invagination (from CE M77-1, Virginia Key, Florida); B, 
Lateral view of spermatheca in V, foreshortened, whole mounted specimen (from CE M77-2, Virginia 
Key, Florida); C, Sectioned spermatheca in V (from CE M77-1); D, Sectioned spermatheca showing 
distinct ampulla and diverticulum, with ectal duct communicating with the ampulla at the origin of 
the diverticulum (from Enchytraeus barkudensis, BMNH 1933.5.25.356, Madras, India). Other ab- 
breviations: a, spermathecal ampulla; d, ampullar diverticulum, extending dorsad; ed, ectal duct; ep, 
circular muscle (black) and epithelial cell layer covered by cuticle; Im, longitudinal muscle; sp, ectal 
spermathecal pore. 


Stephensoniella trevori (Coates, 1980), new combination 
Fig. 3 


Marionina trevori Coates, 1980:1311—1313.—Coates and Ellis, 1981:2137. 
Lumbricillus sterreri Lasserre and Erséus, 1976:453—454 (partim). 


Material examined.—Holotype: NMCIC 1979-1599, whole mount, Qualicum 
Beach, British Columbia, Canada, collected by K.A. Coates, 10 June 1976. Para- 
types: NMCIC 1979-1600; and USNM 58913, whole mounts, Qualicum Beach, 
British Columbia, Canada, collected by K. A. Coates, 10 June 1976. Other ma- 
terial: AH 194, a paratype of Lumbricillus sterreri, whole mount, Shelly Bay, 
Bermuda, collected by P. Lasserre, 1973; CE, 10 whole mounted specimens, just 
north of Cabo Frio, coast of Brazil, collected by J. Renaud-Mornant, 15 Oct. 1976; 
and EVS, 56 whole mounted specimens, Persian Gulf, Saudi Arabia, collected 
by Saudi Arabian Tetra Tech Inc., 1981-1982. 

Description.—Setae slightly sigmoid, to 74 wm long; infrequently with 3 setae 
in postclitellar bundles. Clitellum incomplete medioventrally between male pores. 
Septa 8/9 and 9/10 thickened. Seminal vesicle extending as far anteriad as pos- 


VOLUME 96, NUMBER 3 417 


marina (Moore) sterreri trevori 
as marina as_barkudensis (Lasserre & (Coates) 
Erseus) 
no. of segments S9)= (2) 46-48, 64-67 39-47 35-50 
dorsal vessel origin XIV- XXIII XIl- XXII XIl- XIll XIl>XIll 
sperm funnel L:D 2-2) | 2-38 2-4: 1 (28 || 
sperm funnel collar wide equal funnel equal funnel 
thickened septa (6/7) 7/8-S/O = 7/8- 9/IO 7/8 - Wi0 8/9 - 9/IO 


setae 


spermathecae 


Fig. 3. Distinguishing characteristics of three species of Stephensoniella. Scale bar equals 50 wm. 
Figure of spermatheca of S. sterreri modified after Lasserre and Erséus (1976: fig. 3C). 


terior thickened septum. Vas deferens 10 to 14 wm wide, regular throughout 
length, 2 to 3 times as long as sperm funnel; exiting via simple pore lateral to 
small penial bulb. Spermathecal ampulla and subapical diverticulum approxi- 
mately globular, both 41 to 65 um in diameter; spermathecal ectal duct long and 
uniformly narrow, 11 to 17 wm wide by 36 to 56 wm long; with glands originating 
only around ectal pore. 

Remarks.—Other characteristics are as described for the genus or as previously 
described (Coates 1980; Coates and Ellis 1981). Stephensoniella trevori is distin- 
guished from §. marina (Fig. 3) by size and spermathecal characteristics, as 
discussed for S. sterreri. It is distinguished from the latter species by the distri- 
bution of gland cells only around the pore of the ectal duct of the spermatheca 
and by the relative dimensions of the duct, ampulla and diverticulum of the 
spermatheca (Fig. 3). 

Spermathecal characteristics have been relied upon quite heavily to distinguish 
the two smaller species of Stephensoniella; however, the uniformity of sper- 
mathecal structure observed between and within widely separated populations of 
both S. marina and S. trevori provides a good basis for this reliance. 

The whole mounted specimen of S. trevori from Bermuda (paratype AH 194 
of Lumbricillus sterreri) illustrated by Lasserre and Erséus (1976: Plate 1B) is 0.2 
mm wide, not 0.5 mm as would be determined from the illustration. The extension 
posteriad of the seminal vesicles previously noted (Coates 1980) may be an ar- 


418 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


tefact of fixation or simply related to the volume of the seminal vesicles which 
do not extend anterior to the most posterior thickened septum. 
Habitat.—Upper or mid-intertidal in sand with organic debris and finer sedi- 
ments. 
Distribution.—Qualicum Beach, British Columbia, Canada; Shelly Bay, Ber- 
muda; Persian Gulf, Saudi Arabia; and Brazil. 


Discussion 


When Stephensoniella was first diagnosed (Cernosvitov 1934) it was recognized 
as phylogenetically close to Lumbricillus and Marionina. More recently (Nielsen 
and Christensen 1959) the structural diversity in Marionina has been allowed 
broader limits and the species of Stephensoniella could be included in that genus. 
Stephensoniella, however, combines at least two character states that are uncom- 
mon in Marionina, a nephridium with only the funnel anterior to the septum and 
paired, voluminous seminal vesicles. Distinct, large spermathecal diverticula, seen 
in all of S. marina, S. trevori, and S. sterreri, are unknown in both Marionina 
and Lumbricillus. Thickened, muscular, preclitellar septa are also distinctive for 
the three Stephensoniella species. It would not be surprising if other species 
presently classified, with reservations, as Marionina or Lumbricillus were found 
to belong in Stephensoniella. 


Acknowledgments 


I am indebted to Mr. E. G. Easton, Mr. R. Sims, Dr. J. Renaud-Mornant, Ms. 
J. A. Fournier, Ms. R. Carson, and Dr. C. Erséus for loans of material and for 
providing other useful information. Ms. D. Kathman of EVS Consultants, Sidney, 
B.C., made the Arabian material collected for Saudi Arabian Tetra Tech Inc. 
available to me. Dr. R. O. Brinkhurst and H. R. Baker discussed this work with 
me and provided useful criticisms of the manuscript. Material support was made 
available by Dr. R. O. Brinkhurst, and the University of Victoria provided financial 
support through a Graduate Fellowship. 


Literature Cited 


Cernosvitov, L. 1934. Zur Kenntnis der Enchytraeiden. I—Zoologischer Anzeiger 105: 233-247. 

1935. Oligochaten aus dem Tropischen Stid-Amerika.—Capita Zoologica 6: 1—36. 

——. 1937. System der Enchytraeiden.—Bulletin Association Russe des Recherches Scientifique, 
Prague 5:263—295. 

Coates, K. A. 1980. New marine species of Marionina and Enchytraeus (Oligochaeta, Enchytraei- 

dae) from British Columbia.—Canadian Journal of Zoology 58:1306—1317. 

, and D. V. Ellis. 1981. Taxonomy and distribution of marine Enchytraeidae (Oligochaeta) 

in British Columbia.—Canadian Journal of Zoology 59:2129-2150. 

Kossmagk-Stephan, K. 1983. Marine Oligochaeta from a sandy beach of the Island of Sylt (North 
Sea) with descriptions of four new enchytraeid species.—Mikrofauna Meeresboden (In press). 

Lasserre, P., and C. Erséus. 1976. Oligochetes marins des Bermudes. Nouvelles especes et re- 
marques sur la distribution géographique de quelques Tubificidae et Enchytraeidae.—Cahiers 
de Biologie Marine 17:447—462. 

Moore, J. P. 1902. Some Bermuda Oligochaeta, with a description of a new species.—Proceedings 
of the Academy of Natural Sciences of Philadelphia 54:80-84. 

Nielsen, C. O., and B. Christensen. 1959. Studies on Enchytraeidae. Critical revision and taxonomy 
of European species.—Natura Jutlandica 8—9: 1-160. 


VOLUME 96, NUMBER 3 419 


Stephenson, J. 1915a. On some Indian Oligochaeta from Southern India and Ceylon.—Memoirs of 
the Indian Museum 6:38—108. 

——. 1915b. Oligochaeta, in fauna of Chilka Lake.—Memoirs of the Indian Museum 5:139-146. 

——. 1923. Oligochaeta. Enchytraeidae. Jn The fauna of British India including Ceylon and 
Burma. Edited by Sir A. E. Shipley.—Taylor and Francis, London. Pp. 110-116. 

———. 1930. The Oligochaeta.—Clarendon Press, Oxford, England. 


Department of Biology, University of Victoria, P. O. Box 1700, Victoria, Brit- 
ish Columbia V8W 2Y2, Canada; and Aquatic Zoology Division, British Columbia 
Provincial Museum, Victoria, British Columbia V8V 1X4, Canada. 


PROC. BIOL. SOC. WASH. 
96(3), 1983, pp. 420-428 


DISTOCAMBARUS (DECAPODA: CAMBARIDAE) 
ELEVATED TO GENERIC RANK, WITH AN ACCOUNT OF 
D. CROCKERI, NEW SPECIES, FROM 
SOUTH CAROLINA 


Horton H. Hobbs, Jr., and Paul H. Carlson 


Abstract.—The previously monotypic subgenus Distocambarus proposed by 
Hobbs, 1981, is elevated to generic status, and a new species, Distocambarus 
crockeri, is described from the Savannah River basin of the Piedmont Province 
of South Carolina. The genus now embraces two species: Distocambarus devexus 
(Hobbs, 1981) of the Savannah piedmont of Georgia and D. crockeri which fre- 
quents the piedmont section of the same river basin in South Carolina. Both 
members of the genus are primary burrowers. 


In describing Procambarus (Distocambarus) devexus “‘from the Piedmont 
Province in the Broad River basin (Savannah River watershed) of Oglethorpe and 
Wilkes counties, Georgia,’’ Hobbs (1981:306) chose the subgeneric name to de- 
note his conviction of the remoteness of this crayfish from all known members 
of the genus. He pointed out its similarities to members of the subgenera Gir- 
ardiella, Capillicambarus, Leconticambarus, and Villalobosus and to ‘‘certain 
members of the genus Fallicambarus,’ but he emphasized its distinctive features. 
With the discovery of a close ally of this disjunct species occupying the Savannah 
basin in the Piedmont Province of South Carolina, we believe that these two 
crayfishes, exhibiting such a combination of unique and distinctive characters 
(see the discussion of ‘‘Relationships’’ below), should be recognized at the ge- 
neric level. Therefore we propose that the subgenus Distocambarus be elevated 
to generic rank, and that it encompass Distocambarus devexus (Hobbs) and the 
new species described herein. 


Genus Distocambarus 


Subgenus Distocambarus Hobbs, 1981:301 [Type-species, Procambarus (Disto- 
cambarus) devexus Hobbs, 1981:302]. 


Diagnosis.—Antennal flagellum never with conspicuous fringe on mesial bor- 
der. Third maxilliped with teeth on mesial margin of ischium. Mesial margin of 
palm of chela with row of as many as 8 tubercles; lateral margin of fixed finger 
never bearing spiniform tubercles; opposable margin of dactyl with shallow ex- 
cision proximally. Areola 8 to almost 40 times as long as broad. Ischium of third 
pereiopod only with hook. Coxa of fourth pereiopod lacking caudomesial boss. 
First pleopods of first form male symmetrical, widely separated at base, bearing 
prominent caudoproximal lobe, flexed caudally slightly distal to midlength, and 
partly concealed by sternal setae extending from ventrolateral part of sternum 
and coxae of third, fourth, and fifth pereiopods; terminal elements consisting of 
subtriangular to subconical mesial process, directed caudally to caudodistally, 


VOLUME 96, NUMBER 3 421 


and similarly disposed platelike to bladelike central projection; cephalic process, 
if present, represented by small rounded to subacute knob on cephalodistal end 
of appendage; sternite corresponding to fourth pereiopod conspicuously produced 
ventromesially. Female with annulus ventralis hinged anterodorsally and moving 
through arc of 30 to 90 degrees; sternal plate immediately anterior to annulus 
with narrow median fissure; first pleopod represented by inconspicuous tuber- 
culiform swelling. Branchial count 17 + epipodite. 


Distocambarus crockeri, new species 


Diagnosis.—Body and eyes pigmented, latter small but well developed. Ros- 
trum without marginal spines, tubercles, and median carina. Carapace with one 
to several small cervical tubercles. Areola 7.3 to 13.9 (average 10.2) times as long 
as broad, and constituting 37.6 to 40.9 (average 39.0) percent of entire length of 
carapace (42.7 to 48.4, average 45.3, percent of postorbital carapace length). 
Ventral surface of ischium of third maxilliped only partly obscured by plumose 
setae. First 3 pairs of pereiopods without conspicuous brush of setae extending 
from basis to merus. First pair of pereiopods with ventral surface of merus dense- 
ly tuberculate and corresponding surface of proximal part of both fingers tuber- 
culate. Second pair of pereiopods with conspicuous brush of setae on carpus and 
propodus. First form male with simple hook on ischium of third pereiopod only; 
coxa of fourth pereiopod lacking caudomesial boss. First pleopods widely spaced 
at base, symmetrical, reaching coxae of third pereiopods, with proximomedian 
lobe but without proximomesial spur; cephalic surface with weak shoulder near 
bases of terminal elements; subapical setae absent; shaft of appendage bent cau- 
dodistally near midlength at angle of approximately 40 degrees; terminal elements 
restricted to slender, tapering, distally directed mesial process, and short, 
subquadrate, platelike, corneous central projection directed caudodistally and 
rather strongly mesially; cephalic process absent. Mesial ramus of uropod with 
small distomedian spine premarginal. Female with annulus ventralis capable of 
arclike motion in longitudinal axis of body; large postannular sclerite abutting but 
not covering part of annulus; first pleopods consisting of rudiment in form of 
small tuberculiform protrusion from sternum. 

Holotypic male, form I: Cephalothorax (Fig. la, 1) subovate, compressed lat- 
erally; maximum width of carapace slightly greater than height at caudodorsal 
margin of cervical groove (15.2 and 14.5 mm). Abdomen narrower than thorax 
(12.3 and 15.2 mm). Areola 10.0 times as long as wide with 2 rows of punctations 
across narrowest part. Cephalic section of carapace about 1.6 times as long as 
areola, latter comprising 38.3 percent of total length of carapace (45.8 percent of 
postorbital carapace length). Surface of carapace distinctly punctate dorsally be- 
coming weakly granulate laterally, tubercles slightly larger in hepatic region than 
on most of branchiostegite but largest in anteroventral branchiostegal area. Ros- 
trum broad, gently rounded apically with short triangular acumen reaching mid- 
length of distal podomere of antennular peduncle; margins not thickened; upper 
surface shallowly excavate with usual submarginal punctations and scattered 
moderately large ones. Subrostral ridge clearly defined. Suborbital angle subacute 
and rather prominent. Postorbital ridge moderately strong, ending somewhat 
abruptly anteriorly, neither spine nor tubercle present. Branchiostegal spine small 


422 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 1. Distocambarus crockeri (all illustrations from holotype except c, e from morphotype, and 
k from allotype): a, Lateral view of carapace; b, c, Mesial view of first pleopod; d, Caudal view of 
first pleopods; e, f, Lateral view of first pleopod; g, Antennal scale; h, Epistome; i, Dorsal view of 


carapace; j, Proximal podomeres of third, fourth, and fifth pereiopods; k, Annulus ventralis; 1, Dorsal 
view of distal podomeres of cheliped. 


VOLUME 96, NUMBER 3 423 


but acute; cervical spine absent but row of small tubercles along posterior side 
of region of junction of anterior and posterior arms of cervical groove. 

Abdomen shorter than carapace (26.6 and 31.3 mm). Pleura of second through 
fifth abdominal segments broadly rounded to subtruncate ventrally, lacking cau- 
doventral angles. Cephalic section of telson with 2 spines (more mesial one mov- 
able) in each caudolateral corner. Cephalic lobe of epistome (Fig. 1h) triangular, 
anterolateral margins elevated ventrally; main body of epistome with anterome- 
dian depression but lacking fovea; epistomal zygoma broadly arched. Ventral 
surface of proximal podomere of antennular peduncle with very small spine dis- 
tinctly distal to midlength. Antennal peduncle without tubercles and spines on 
basis and ischium; flagellum with distal articles lacking but reaching second ab- 
dominal tergum. Antennal scale (Fig. 1g) about 2.1 times as long as broad, widest 
distal to midlength; greatest width of lamellar area almost twice that of thickened 
lateral part. 

Third maxilliped extending anteriorly beyond antennal peduncle by length of 
dactyl and half that of propodus; mesial sector of ventral surface of ischium 
densely clothed in long plumose setae; lateral sector with closely spaced shorter 
ones proximally and yet shorter, more widely placed ones distally; distolateral 
margin of article produced in short spine; merus with setae similarly disposed. 

Right chela (left probably regenerated) (Fig. 11) subelliptical in section, some- 
what depressed; mesial margin of palm |.1 times as long as greatest width, former 
about two-fifths that of length of palm; almost entire surface of palm with squa- 
mous to subsquamous tubercles. Mesial margin of palm with row of 8 (right) or 
6 (left) tubercles flanked dorsally by row of 8 (right) or 6 (left) and irregular ventral 
row of 8, few additional tubercles present between rows; ventral surface with 2 
prominent tubercles opposite base of dactyl. Both fingers with moderately well 
defined dorsomedian ridges; that on fixed finger flanked along proximal half by 
tubercies and distally by row of punctations, that on dactyl by tubercles along 
proximal three-fourths and by punctations distally. Opposable margin of fixed 
finger with row of 9 (right) or 7 (left) tubercles, second from base largest, scattered 
along almost entire length of fingers, and single row of minute denticles extending 
between, and some under, tubercles; 2 tubercles also present slightly below row: 
larger one at about midlength and smaller at about base of apical third of finger; 
lateral surface with row of tubercles on proximal third replaced along distal part 
of finger by row of setiferous punctations. Opposable margin of dactyl with row 
of 10 (right) or 11 (left) tubercles, second from base largest, dispersed along almost 
entire length of finger, interrupting row of minute denticles; basal fourth of margin 
shallowly excavate; mesial surface of dactyl tuberculate with row extending from 
base almost to corneous tip of finger, becoming subacute on distal part of pod- 
omere; ventral surface of fingers with tubercles flanking median ridge proximally 
and punctations distally. 

Carpus of cheliped distinctly longer than mesial margin of palm (10.5 and 8.1 
mm), its dorsal surface with broad, slightly sinuous, shallow furrow extending 
from almost 0.2 length of podomere from proximal end to about same distance 
from distal end, and with scattered punctations and few tubercles proximome- 
sially; mesial surface with irregular dorsal row of 7 or 8 small tubercles below 
which 14 or 15 additional tubercles, only one of which decidedly larger than 


424 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


others, also present, and premarginal setal band distomesially; distoventral mar- 
gin of carpus with 2 tubercles, larger, more lateral one bearing articular socket 
receiving ventrolateral condyle of propodus. 

Merus with usual tubercles dorsally, none acute, and more distal ones not 
conspicuously larger than several proximal to it; mesial and lateral surfaces most- 
ly smooth, but distomesial area bearing number of very small tubercles; ventral 
surface studded with crowded tubercles, irregular mesial and lateral rows con- 
sisting of 15 or 16 tubercles, spiniform. Ischium with row of 4 small tubercles 
mesially. 

Hook on ischium of third pereiopod (Fig. 1j) simple, comparatively heavy, and 
overreaching basioischial articulation, latter not opposed by tubercle on basis. 
Coxa of neither fourth nor fifth pereiopods with boss; ventral membrane of coxa 
of fifth conspicuously setose. 

Sternum between third pereiopods rather shallow; that between both fourth 
and fifth comparatively deep; lateral part of that between fourth produced ven- 
trally in moderately strong, posteroventrally projecting lobe. Plumose pubescence 
associated with sternum and coxae of all pereiopods very prominent. 

First pleopods (Fig. Ib, d, f) as described in ‘‘Diagnosis.’’ Uropods with both 
lobes of proximal podomere bearing acute corneous spines; mesial ramus with 
distomedian spine small and situated premarginally. 

Allotypic female: Differing from holotype, other than in secondary sexual fea- 
tures, in following respects: areola 9.6 times as long as broad; postcervical groove, 
which not evident in holotype, clearly defined and situated 0.2 areola length 
posterior to cervical groove; tubercles on carapace much weaker; rostrum almost 
reaching distal margin of ultimate podomere of antennular peduncle. Suborbital 
angle weak and obtuse; branchiostegal spine smaller but distinct; only 1 or 2 
cervical tubercles present; third maxilliped only slightly overreaching antennal 
peduncle. Length of mesial margin of palm of chela equal to width and about 0.4 
as long as chela; only | tubercle (injured) on ventral surface at base of dactyl; 
opposable margin of fixed finger with only | tubercle below principal row; 7 
tubercles on corresponding margin of dactyl, and more mesial of 2 tubercles on 
distoventral margin of carpus less conspicuous than in holotype. (See measure- 
ments in Table 1.) 

Annulus ventralis (Fig. 1k) 1.7 times as wide as long, hinged cephalically, 
moving through arc of 90 degrees, D-shaped with anterior margin almost straight 
and slightly elevated; ventral surface sloping posteriorly and mesially from an- 
terolateral angles; sinus almost straight, beginning at about midlength of annulus, 
slightly dextral to median line, and extending caudosinistrally, ending on cau- 
dosinistral wall of annulus; sinistral wall of sinus increasing in height posteriorly, 
forming conspicuous prominence on caudal wall. Postannular sclerite subtrape- 
zoidal in shape, its maximum width 1.5 times its length; sclerite 0.9 as wide and 
0.9 as long as annulus, its anteroventral margin with symmetrical pair of broad 
excavations rendering margin with 3 short rounded projections. 

Sternum anterior to annulus with narrow, median, longitudinal cleft. First pleo- 
pod represented by tuberculiform rudiment. 

Morphotypic male, form II: Differing from holotype, other than in development 
of secondary sexual characters, in only few minor respects: epistome, while sub- 
triangular, with more irregular anterolateral margins; mesial margin of palm of 


VOLUME 96, NUMBER 3 425 


Table 1.—Measurements (mm) of Distocambarus crockeri. 


Holotype Allotype Morphotype 

Carapace 

Entire length 31.3 34.6 23.8 

Postorbital length 26.2 29.5 20.4 

Width 1572 16.2 11.1 

Height 14.5 15.0 10.5 
Areola 

Width 1.2 1.4 1.1 

Length 12.0 13.5 
Rostrum 

Width 5.0 Soll 4. 

Length 6.8 7.0 5)o 
Right chela 

Length, palm mesial margin 8.1 9.2 4.9 

Palm width 7.6 9.2 Do 

Length, lateral margin 20.8 21.4 12.5 

Dactyl length 6.9 12.7 V2 
Abdomen 

Width 12.3 13.8 9.0 

Length 26.6 31.8 23.3 
Carpus of cheliped 

Width Sod 6,2 3.7 

Length 10.5 10.8 6.4 


chela with row of 8 or 9 tubercles; opposable margin of fixed finger with row of 
3 or 4 tubercles along proximal half, and corresponding margin of dactyl with 2 
tubercles representing proximal 2 in holotype. (See measurements in Table 1.) 

Hook on ischium of third pereiopod much less conspicuous and not projecting 
over basioischial articulation. First pleopod (Fig. Ic, e) not distinctly reflexed; 
mesial process longer and distal part more slender than in holotype; central pro- 
jection rounded and non-corneous but disposed as in holotype; juvenile suture 
evident. 

Color notes.—(Based upon specimens from burrows at junction of U.S. High- 
way 378 and County Road 423, northwest of Edgefield, Edgefield County, South 
Carolina.) Ground color shades of brown. Dorsum of cephalic region and areola 
dark brown, rostral margins and V-shaped marking in posterior gastric region 
almost black; areola flanked by grayish brown longitudinal stripes which flanked 
laterally by dark chocolate stripes extending from cervical groove to posterior 
margin of carapace; lateral surface of branchiostegites with irregular tan splotches 
on brown fading ventrally from just-mentioned chocolate stripe to pale brown 
along ventral margin. Abdomen with orange tan dorsomedian stripe flanked by 
broad dark chocolate stripes (continuous with those on carapace) extending from 
first to anterior part of sixth segment; terga ventrolateral to chocolate stripe tan 
with reticulate darker brown mottlings; pleura, except for anteroventral tan sec- 
tions, bearing reticulate pattern of dark brown; tergum of sixth segment, telson, 
and uropods with brown mottlings on tan. Cheliped basically dark tan with brown 
tubercles; very dark brown markings on distal margin of merus, dorsolateral 


426 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


surface of carpus, dorsomesial part of palm, and dorsal part of fingers; tips of 
latter reddish orange; remaining pereiopods with broad irregular bands, darker 
at distal ends of podomeres; merus and carpus each with proximal and distal 
bands. 

Type-locality.—Roadside ditch 0.7 miles south of Parksville, McCormick Coun- 
ty, South Carolina, on U.S. Highway 221. The ditch from which the specimens 
were dug was partly shaded by trees of the genera Pinus, Juniperus, and Acer 
and had been scraped in making road repairs. Water stood in the lower section, 
and many burrows that were not inundated were surrounded by pellets composed 
of sandy clay that recently had been brought to the surface. The burrows were 
comparatively simple: none that were excavated had more than three openings, 
and although branching, possessed only one deep passage. Some of the burrows 
contained several individuals, but whether or not the multiple occupancy resulted 
from the young-of-the-year not having left the parent burrow could not be deter- 
mined. Burrows elsewhere containing more than one individual housed only a 
first form male and a female, or a female with few to several juveniles. No 
other crayfish species was found in the colony at this locality. 

Disposition of types.—The holotypic male, form I, the allotype, and the mor- 
photypic male, form II, are deposited in the National Museum of Natural History 
(Smithsonian Institution), nos. 178582, 178583, and 178584, respectively, as are 
the paratypes consisting of 7 males, form I, 2 males, form II, 15 females, 7 juvenile 
males and 7 juvenile females. 

Size.—The largest specimen available is a first form male having a carapace 
length of 34.7 mm (postorbital carapace length 30.0 mm); corresponding lengths 
of the smallest first form male are 27.8 (24.1) mm, and those of the largest female, 
the allotype, 34.6 (29.5) mm. Ovigerous females have not been found. 

Range and specimens examined.—This crayfish appears to be restricted to the 
eastern watershed of the Savannah River in the Piedmont Province of South 
Carolina. Specimens have been examined from the following localities: Mc- 
CORMICK COUNTY—(1) type-locality, | dI, 1 SII, 4 2,4jd6, 552, 19 Apr 
1981, G. B. Hobbs, J. E. Pugh, HHH; (2) roadside seepage area 0.9 mi S of 
Abbeville Co line on Co Rd 81, 1 dI, 1 2, 1 jd, 18 Apr 1981, GBH, JEP, HHH; 
(3) roadside ditch 0.5 mi E of Stephens Cr on St Rte 283, 2 2, 1 jd, 24 Apr 1982, 
GBH, HHH. ABBEVILLE COUNT Y—(4) seepage and wet area around farm 
pond 2.6 mi NE of Calhoun Falls on St Rte 72, 2 dI, 17 Apr 1981, JEP, HHH; 
(5) roadside ditch 100 yds S of Gill Creek on Co Rd 32, 1 oI, 1 2, 25 Apr 1982, 
GBH, HHH. EDGEFIELD COUNT Y—(6) roadside ditch at westernmost trib- 
utary to Rocky Creek at U.S. Hwy 378, 2 mi W of Co Rd 51, 1 oI, 1 oll, 4 2, 
1j3d,2j2, 9 Apr 1982, P. H. Carlson and E. M. Younginer; (7) roadside ditch 
at jct of U.S. Hwy 378 and Co Rd 423, NW of Edgefield, 2 oI, 1 dII, 4 9, 24 
Apr 1982, GBH, HHH. (Burrows of Cambarus (D.) latimanus (LeConte) were 
excavated within a meter of that of one of these specimens.) 

Variations.—Although there are many minor variations among the specimens 
that have been examined, none involves the differences between this and closely 
or distantly related species. The rostral margins are often more strongly conver- 
gent than that illustrated, and the carpus of the cheliped may not be so narrow 
as that reported for the holotype, but never are there massive tubercles on the 


VOLUME 96, NUMBER 3 427 


mesial surface of the latter; always the carpus appears abnormally long, and 
though tuberculate, the mesial surface lacks the conspicuously dominant major 
spine that is typical of most crayfishes in the eastern part of the United States. 
The only difference noted in the first pleopods of the first form male in specimens 
from the few localities from which they are available is in the degree of sclero- 
tization of the central projection, but none of the variations seems to be correlated 
with specimens from a restricted part of the range of the species. 

Relationships.—Distocambarus crockeri is more closely allied to D. devexus 
than to any other crayfish. Except for references to the secondary sexual features, 
most of the description of the latter (Hobbs 1981:302) applies quite well to D. 
crockeri. The only striking differences between the two exist in features of the 
first pleopod of the male and those of the annulus ventralis. Ecologically, one 
appears to vicariate for the other on opposite sides of the Savannah River. Where- 
as their broad areolae and ungainly legs suggest a better adaptation to life in 
surface lentic or lotic habitats, both are primary burrowers. Their relationships 
to other members of the Cambarinae are not understood, but perhaps the kinships 
suggested by Hobbs (1981:43, and fig. 11) are not totally erroneous. 

These two crayfishes may be distinguished from all others by the following 
combination of characters: rostrum without marginal spines; ventral surface of 
merus of cheliped densely tuberculate; ischia of only third pereiopods with hooks; 
coxae of fourth pereiopods without caudomesial boss; first pleopods of first form 
male symmetrical, widely separated basally, with prominent caudoproximal lobe, 
deflected caudodistally near midlength, and lacking caudal process; carpus of 
cheliped twice as long as wide; annulus ventralis moving through arc of approx- 
imately 90 degrees; postannular sclerite large and platelike; and sternum of female 
anterior to annulus narrowly cleft. 

Distocambarus crockeri may be separated from D. devexus by the first pleopod 
of the male, which lacks a cephalic process and possesses a more acute mesial 
process and a much larger quadrangular central projection, and by the annulus 
ventralis which when depressed is not partly overlapped by the postannular 
sclerite. 

Ecological notes.—The burrows of Distocambarus crockeri do not differ in any 
obvious way from those of D. devexus, which Hobbs (1981:307) described as 
‘‘moderately complex.’’ Most of those excavated had two or three passageways 
leading to the surface, at least one topped by a crude turret; these tunnels con- 
verged to form one subvertical gallery that penetrated the water table and oc- 
casionally branched into two or three passages directed downward; all ended 
blindly, in some instances more than a meter below the surface. Burrows were 
found in seepage areas, bogs, and in both waterlogged and comparatively dry 
roadside ditches. In all except one of the localities, clumps of a sedge, 0.3 to 0.6 
m in height, were among the most conspicuous plants present. In fact, the 
presence of the sedge led one of us (H.H.H.) to choose collecting sites where 
this crayfish was found as he was driving along the county roads and highways. 
(See “‘Type-locality.’’) 

Etymology.—This crayfish is named in honor of Denton W. Crocker, a fellow 
student of crayfishes and a friend, whose contributions to our knowledge of the 
American crayfish fauna are invaluable. 


428 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Acknowledgments 


We extend our thanks to Georgia B. Hobbs and Jean E. Pugh, Christopher 
Newport College, both long time collecting companions, and to Edward M. Youn- 
giner, South Carolina Department of Health and Environmental Control, for their 
help in obtaining the specimens on which this description is based, and to Ray- 
mond W. Bouchard, Wildwood Crest, New Jersey, and Raymond B. Manning of 
the Smithsonian Institution, for their criticisms of the manuscript. 


Literature Cited 


Hobbs, Horton H., Jr. 1981. The crayfishes of Georgia.—Smithsonian Contributions to Zoology, 
318: vi + 549 pages, 262 figures. 


(HHH) Department of Invertebrate Zoology, Smithsonian Institution, Wash- 
ington, D.C. 20560; (PHC) Department of Health and Environmental Control, 
2600 Bull Street, Columbia, South Carolina 29201. 


PROC. BIOL. SOC. WASH. 
96(3), 1983, pp. 429-439 


DISTOCAMBARUS (FITZCAMBARUS) CARLSONI, 
A NEW SUBGENUS AND SPECIES OF CRAYFISH 
(DECAPODA: CAMBARIDAE) FROM 
SOUTH CAROLINA 


Horton H. Hobbs, Jr. 


Abstract.—A new subgenus, Fitzcambarus, is proposed to receive Distocam- 
barus (Fitzcambarus) carlsoni, the third species assigned to the genus. This cray- 
fish, differing from its relatives in possessing a comparatively narrow areola, has 
been found in a single locality in the Saluda River basin of Anderson County, 
South Carolina. Like its congeners, it is a primary burrower and shares a swamp- 
seepage area with the crayfish Cambarus (Jugicambarus) carolinus (Erichson), 
the only primary burrower previously reported from the basin. 


The crayfish described here was discovered in burrows constructed in a swampy 
area bordering a tributary to the Saluda River in Anderson County, South Car- 
olina. Late in the afternoon of 27 April 1982, after having collected crayfishes in 
a small sand and silt bottomed stream, I discovered burrows in the adjoining 
swamp. The only burrowing crayfish that I had encountered previously in the 
upper Saluda Basin was Cambarus (Jugicambarus) carolinus (Erichson, 1846), 
and I anticipated that this was the crayfish responsible for the piles of earth 
scattered over the swamp floor. Because of the densely matted roots just beneath 
the surface of the ground, attempts to excavate two of the burrows resulted in 
failure to obtain an adult individual, but a very small juvenile was retrieved from 
one of them. The specimen was brought into the laboratory where it was found 
to belong to an unknown species. My friend Paul H. Carlson was advised of the 
find, and he visited the locality in May when he obtained several representatives 
of C. (J.) carolinus. Another attempt by him to obtain additional specimens of 
the new crayfish was made in June when he succeeded in securing several spec- 
imens; however, not until early in November, when both of us spent the better 
part of seven hours digging in the swamp, was an adequate series, including two 
first form males obtained by Dr. Carlson, acquired for preparing this description. 

This crayfish is named in honor of Dr. Carlson, of the South Carolina Depart- 
ment of Health and Environmental Control, in token of his indefatigable efforts 
in sampling so many burrowing crayfish populations occurring in South Carolina. 

Reasons for proposing the recognition of subgenera in the genus Distocambarus 
are pointed out in the discussion of ““Relationships’’ below. Diagnoses and the 
compositions of the subgenera are as follows: 


Genus Distocambarus Hobbs, 1981 
Subgenus Distocambarus 


Diagnosis.—Areola no more than 14 times as long as wide; mesial margin of 
palm of chela of male longer than maximum width of podomere and at least as 
long as carpus; terminal elements of first pleopod of first form male directed 


430 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


caudodistally, central projection platelike and comparatively short; annulus ven- 
tralis movable through arc of 90 degrees, and postannular sclerite large, subtrape- 
zoidal, and almost as long as annulus. 

Species.—Distocambarus (Distocambarus) devexus (Hobbs, 1981:302) and 
Distocambarus (Distocambarus) crockeri Hobbs and Carlson, 1983:421. 


Fitzcambarus, new subgenus 


Diagnosis.—Areola at least 15 times as long as wide; mesial margin of palm of 
chela of male shorter than wide and shorter than carpus; terminal elements of 
first pleopod of first form male directed caudally, central projection bladelike and 
comparatively long; annulus ventralis movable through arc of less than 50 de- 
grees, and postannular sclerite small, subovate in outline, wedge-shaped, and 
distinctly shorter than annulus. 

Gender.—Masculine. 

Type-species.—Distocambarus (Fitzcambarus) carlsoni new species. 

Etymology.—This name is derived from the surname Fitzpatrick and the cray- 
fish genus Cambarus. | am pleased to propose this taxon honoring Joseph F. 
Fitzpatrick, Jr., a friend and an able student of crayfishes who has added much 
to our knowledge of the Cambaridae. 


Distocambarus (Fitzcambarus) carlsoni, new species 
Fig. | 


Diagnosis.—In addition to characters cited for genus, rostrum broadly acu- 
minate, margins little thickened and devoid of spines or tubercles. Cephalic ex- 
tremity of postrobital ridge merging almost imperceptibly with carapace. Subor- 
bital angle obtuse and inconspicuous. Cervical spine lacking. Areola 15.2 to 38 
times as long as wide, comprising 38.1 to 41.1 percent of entire length of carapace 
(43.7 to 47.3 percent of postorbital carapace length) and never with more than 1, 
often none, punctation in narrowest part. Chela with well defined row of 5 to 7 
tubercles on mesial surface of palm, additional row flanking it dorsolaterally and 
2 or 3 tubercles distoventrally. No tubercle on basis of third pereiopod opposing 
hook on ischium. 

Holotypic male, form I: Cephalothorax (Fig. la, m) subcylindrical; maximum 
width of carapace slightly greater than height at caudodorsal margin of cervical 
groove (13.2 and 12.3 mm). Abdomen distinctly narrower than thorax (9.6 and 
13.2 mm). Areola 38.3 times as long as wide, with room for no more than 1 
punctation across narrowest part. Cephalic section of carapace about 1.4 times 
as long as areola, latter comprising 41.1 percent of total length of carapace (47.3 
percent of postorbital carapace length). Surface of carapace mostly punctate, few 
tubercles present in hepatic region and in anteroventral branchiostegal region. 
Rostrum broad and short, gently rounded apically with small, poorly delimited 
acumen reaching base of distal podomere of antennular peduncle; margins not 
conspicuously thickened; upper surface shallowly concave with usual submar- 
ginal punctations and moderately large ones posteriorly. Subrostral ridge weak 
but clearly defined. Suborbital angle broadly obtuse, very weak. Postorbital ridge 
moderately well developed but merging insensibly with carapace cephalically. 
Branchiostegal and cervical spines absent. 


VOLUME 96, NUMBER 3 431 


Fig. 1. Distocambarus (F.) carlsoni (all from holotype except c, e from morphotype, i-—/ from 
paratypic female, and n from allotype): a, Lateral view of carapace; b, c, Mesial view of first pleopod; 
d, Caudal view of first pleopods; e, f, Lateral view of first pleopod; g, Epistome; 4, Proximal pod- 
omeres of third, fourth, and fifth pereiopods; 7, Antennal scale; /, Incisor lobe of mandible; k, /, Molar 
lobe of mandible; m, Dorsal view of carapace; n, Annulus ventralis and adjacent sternites; 0, Denuded 
ventrolateral parts of sternum and ventral articular regions of left third and fourth pereiopods showing 
distinctly produced posteroventral lobe on latter; p, Dorsal view of distal podomeres of cheliped. 


432 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Abdomen shorter than carapace (22.8 and 28.0 mm). Pleura of second through 
fifth abdominal segments rounded ventrally, lacking caudoventral angle. Cephalic 
section of scabrous telson with 2 spines (more mesial one movable) in each cau- 
dolateral corner. Cephalic lobe of epistome (Fig. 1g) subtrapezoidal, cephalic part 
truncate with small asymmetrical median projection, margins weakly elevated 
ventrally; main body of epistome with triangular median depression but lacking 
distinct fovea; epistomal zygoma broadly arched. Ventral surface of proximal 
podomere of antennular peduncle with heavy short spine distal to midlength. 
Antennal peduncle without tubercles and spines on basis and ischium; flagellum 
with distalmost articles lacking, but almost reaching midcaudal margin of cara- 
pace. Antennal scale (Fig. li) about 2.5 times as long as broad, widest slightly 
distal to midlength; greatest width of lamellar area about 1.5 times that of thick- 
ened lateral part. 

Third maxilliped extending as far anteriorly as antennal peduncle; mesial sector 
of ventral surface of ischium with clusters of stiff, long setae some of which 
flanked laterally by shorter, plumose ones; latter also present in lateral sector 
where conspicuous in submarginal lateral row; merus with setae similarly dis- 
posed. 

Right chela (Fig. Ip) subelliptical in section, rather strongly depressed; palm 
1.4 times as broad as length of mesial margin; length of latter one-third that of 
entire length of chela; almost entire surface of palm with squamous to subsqua- 
mous tubercles. Mesial margin of palm with row of 7 tubercles flanked dorsolat- 
erally by row of 4 (right) or 5 (left) and few additional ones. Both fingers with 
well defined dorsomedian ridges flanked by setiferous punctations. Opposable 
margin of fixed finger with row of 3 (left with 4) tubercles, second from base 
largest, along proximal two-fifths of finger and single row of minute denticles 
extending between and distal to tubercles from base of finger to corneous tip; 
acute tubercle also present slightly below row at about midlength of finger; lateral 
margin of finger weakly costate with row of punctations. Opposable margin of 
dactyl with 2 tubercles, more distal larger, on proximal half of finger interrupting 
single row of minute denticles extending from proximal base of proximalmost 
tubercle to corneous tip of finger, basal half of margin shallowly excavate; mesial 
margin of dactyl with row of 3 tubercles proximally followed by series of setif- 
erous punctations, tubercles decreasing in size and becoming more squamous 
distally. 

Carpus of cheliped distinctly longer than mesial margin of palm of chela (8.6 
and 5.8 mm), with shallow sinuous furrow extending almost entire length of pod- 
omere, dorsal surface of which sparsely punctate; mesial surface with dorsal row 
of 5 or 6 small tubercles, below them 6 or 7 additional ones, distalmost of which 
conspicuous; prominent tuft of plumose setae situated distomesially; distoventral 
margin of carpus with 2 tubercles, larger, more lateral one bearing articular socket 
receiving ventrolateral condyle on propodus. 

Merus with usual tubercles dorsally, 3 distal tubercles somewhat larger than 
others; mesial and lateral surfaces rather smooth; ventral surface with crowded 
tubercles, mesial row consisting of 11 tubercles, and less regular lateral row of 
10 or 11. All tubercles on merus comparatively small, and none spiniform. Is- 
chium with | well defined tubercle proximomesially and 2 or 3 vestigial ones 
distal to it. 


VOLUME 96, NUMBER 3 433 


Hook on ischium of third pereiopod (Fig. 1h) simple and acute, overreaching 
basioischial articulation, not opposed by strong tubercle on basis. Coxa of neither 
fourth nor fifth pereiopod with boss; ventral membrane of coxa of fifth conspic- 
uously setose. 

Sternum between third pereiopods rather shallow; that between both fourth 
and fifth comparatively deep, ventrolateral elements corresponding to third and 
fourth produced ventrally much beyond articulation with respective coxae; that 
associated with fourth very strong, subtriangular, and curved ventromesially (Fig. 
lo). Plumose pubescence associated with sternum and coxae of all pereiopods 
very prominent. 

First pleopods (Fig. 1b, d, f) symmetrical, not contiguous at base, reaching 
coxae of third pereiopods, bearing strong caudoproximal lobe and broadly round- 
ed proximomedian lobe, flexed caudally slightly distal to midlength, and lacking 
subapical setae. Terminal elements limited to mesial process and central projec- 
tion, directed caudolaterally and caudomesially, respectively, at about 90 degrees 
to axis of proximal half of appendage; mesial process non-corneous, tapering, 
and reaching almost as far caudally as corneous, bladelike central projection 
which provided with subapical notch. 

Uropods with both lobes of proximal podomere bearing acute spines; mesial 
ramus with distomedian spine small and situated distinctly premarginally. 

Allotypic female: Differing from holotype, other than in secondary sexual fea- 
tures, in following respects: areola 23.4 times as long as wide; cephalic section 
of carapace 1.5 times as long as areola, latter comprising 39.9 percent of total 
length of carapace (45.7 percent of postorbital carapace length); chela propor- 
tionately smaller than that in male, mesial margin of palm with row of 6 (right) 
or 8 (left) tubercles flanked dorsolaterally by row of 6; opposable margin of fixed 
finger with row of 3 tubercles, that of dactyl also with 3, second from base largest; 
mesial margin of dactyl with longitudinal row of 3 (left) or 4 (right) tubercles 
basally, row flanked by 2 tubercles proximoventrally; distal tubercle in ventrome- 
sial row on merus of cheliped subspiniform; 3 tubercles on ventromesial margin 
of ischium of cheliped better developed than those in holotype. (See Table 1.) 

Sternum between fourth pereiopods with very narrow median fissure (expand- 
ing anteriorly and posteriorly) extending entire length. 

Annulus ventralis (Fig. In) hinged cephalically (moving through arc of some 
30 degrees), approximately 1.4 times as broad as long; elevated lateral areas 
separated by sinistrally disposed depression, latter broad anteriorly, gradually 
narrowing posteriorly, and terminating in fossa situated sinistral to median line 
slightly posterior to midlength of annulus; sinus, originating in fossa, coursing 
dextrally across median line before forming broad arc caudosinistrally and ending 
on caudal wall of annulus slightly dextral to median line. 

Postannular sclerite ovate, 0.7 as wide and 0.5 as long as annulus; central area 
elevated (ventrally) with anterior face of elevation plane and posterior surface 
rounded. 

Morphotypic male, form II: Except in secondary sexual characters, differing 
in no conspicuous way from holotype; few differences noted, probably reflecting 
juvenile condition of specimen: anterior section of telson with only 1 spine in 
each caudolateral corner; cheliped with fewer well developed tubercles on merus 
and carpus, and lateral lobe of proximal podomere of uropod without well defined 


434 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Table 1—Measurements (mm) of Distocambarus (F.) carlsoni. 
Holotype Allotype Morphotype 

Carapace 

Entire length 28.0 29.3 21.9 

Postorbital length 24.3 25.6 18.8 

Width 13.2 14.2 9.6 

Height 12.3 12.5 9.0 
Areola 

Width 0.3 0.5 0.4 

Length IES) 11.7 8.3 
Rostrum 

Width 4.6 4.6 3.6 

Length 4.5 4.6 Sol 
Chela 

Length, palm mesial margin 5.8 5.4 3.6 

Palm width 8.2 VES 4.8 

Length, lateral margin 17.6 D)o7/ 11.2 

Dactyl length 10.5 9.6 Holl 
Abdomen 

Width 9.6 10.5 6.9 

Length 22.8 24.5 16.0 
Carpus of cheliped 

Width Dall 5.4 3.6 

Length 8.6 8.5 5.6 


spine. Ischium of third pereiopod with only rudiment of prominent hook in ho- 
lotype, but produced posteroventral lobe of ventrolateral element of sternum 
corresponding to fourth pereiopod clearly evident. First pleopod (Fig. Ic, e) lack- 
ing strong caudal flexure near midlength; central projection non-corneous, and 
both terminal elements bent caudally at about 90 degrees to shaft, but shorter, 
less sharply delimited, and distal parts more inflated than in holotype. 

Color notes.—Dorsum of carapace brown with reddish or greenish suffusion. 
Lateral rostral carinae, postorbital ridges, and small lateral spot on posterior 
margin of cervical groove very dark, almost black. Dark dorsum of thoracic region 
abruptly replaced dorsolaterally by lavender cream extending ventrally over bran- 
chiostegites to very pale marginal carina; anteroventral branchiostegal region 
cream to white. Lateral surface of cephalic section of carapace with large irregular 
brownish area in hepatic region extending anteriorly as narrow stripe across upper 
orbital region, remainder pinkish lavender. Abdomen with first abdominal tergum 
very dark reddish purple, second through fourth segment with rectangular, dark, 
reddish brown markings along median line, decreasing in size posteriorly to sixth 
where sublinear, but expanding and forming triangular marking on anterior sec- 
tion of telson; longitudinal stripe extending along dorsal parts of pleura onto 
anterolateral surface of telson. Anteroventral parts of pleura pale pinkish cream, 
remainder pinkish tan. Ground color of telson and uropods pale tan with pinkish 
suffusion, although margins, sutures, and ridges pale brown. Antennular and 
antennal peduncles reddish brown, flagella tan. Chelipeds basically pinkish lav- 
ender with very dark brown dorsal stripe beginning at base of merus and broad- 


VOLUME 96, NUMBER 3 435 


ening over distal part of podomere; carpus with dark stripes flanking dorsomedian 
depression and splotches around bases of mesial tubercles; propodus with dark 
dorsomesial area studded with very dark tubercles; band across distal part of 
palm extending distolaterally on fixed finger. Dactyl with triangular dark wedge 
on dorsomesial surface, its fading apex approaching distal end of finger. Second 
pereiopod with merus as in cheliped, and carpus with stripe on dorsal margin and 
another laterally, chela pale grayish tan. Remaining pereiopods with dorsal parts 
of podomeres from merus distally dark, intensifying at articular margins. Ventral 
surfaces of all pereiopods pale pinkish lavender, especially chela, to cream. 

Type-locality.—A swampy area bordering an unnamed tributary of the Saluda 
River about | mile north of State Route 81 on Route 106. There, in an area shaded 
by Liriodendron tulipifera, Liquidambar styraciflua, Alnus rugosa, Pinus sp., 
and Quercus sp., the water table fluctuates from the surface to about 0.8 m 
below it. The soil is a sandy clay, in some areas rich in humus, and a layer 
of decaying leaves litters the ground. Chimneys constructed by Distocambarus 
(F.) carlsoni and Cambarus (J.) carolinus (Erichson, 1846) and a few fallen 
branches from the overhanging trees are the only conspicuous irregularities in 
the gently sloping swamp floor. Matted roots of the many trees and shrubs lie at 
and within a few centimeters below the surface. The colony from which the type- 
series was obtained occupies an area of some 200 square meters. (See “*Ecological 
Notes’ below.) 

Disposition of types.—The holotype, allotype, and morphotype are deposited 
in the National Museum of Natural History (Smithsonian Institution), nos. 178599, 
178600, and 178601, respectively, as are the paratypes consisting of 1 male, form 
I, 13 females, 2 juvenile males, 4 juvenile females, and | ovigerous female. 

Size.—The largest specimen available is a female having a carapace length of 
31.0 mm (postorbital carapace length 27.0 mm); corresponding lengths of the 
smaller of the two first form males are 25.8 (22.7) mm, and those of the ovigerous 
female, 27.1 (23.8) mm. 

Range and specimens examined.—This crayfish is known from only the type- 
locality, in the Saluda River basin of South Carolina: Anderson County—(1) type- 
locality, 1 32, 27 Apr 1982, G. B. Hobbs and HHH; | GII, 1 2, | ovig. 
(carapace length 27.1 mm, carrying 25 eggs with diameters of 2.0 to 2.1 mm), | 
jo,2j2, 19 Jun 1982, P. H. Carlson; 3 9, 22 Oct 1982, PHC; 1 2, 1j2, 1 Nov 
1982, GBH, HHH; 2 oI, 9 2, 1 jd and some 20 very small juveniles which are 
being maintained alive, 2 Nov 1982, PHC, GBH, HHH. 

Relationships.—Distocambarus (Fitzcambarus) carlsoni is a unique crayfish 
which, while bearing strong resemblances to members of the genus Cambarus, 
does not share several characteristics that typify all of the now recognized species 
assigned to that genus. The features that are unlike those in any member of 
Cambarus are: 

(1) The bent shaft of the first pleopod of the first form male (Figs. 15, f). In all 
members of Cambarus it is straight, or at most slightly curved, and the terminal 
elements, with few exceptions, project from it at no less than 90 degrees. In D. 
(F.) carlsoni, the shaft is rather strongly reflexed slightly distal to midlength, and 
the terminal elements are disposed caudally at much less than 90 degrees to the 
distal segment of the shaft. 

(2) A strong caudoproximal lobe on the first pleopod of the first form male (Fig. 


436 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


lb, f). In most members of Cambarus there is hardly a trace of such a lobe, and 
in none is it nearly so prominent as it is in D. (F.) carlsoni. 

(3) No boss on the coxa of the fourth pereiopods of the first form male (Fig. 
Ih). In all members of Cambarus, a well defined boss is present on the cau- 
domesial angle of the coxa of this appendage; no such swelling is evident in the 
species described here. 

(4) Produced lateral lobe of the sternum associated with the fourth pereiopods 
of the first form male (Fig. lo). In Cambarus, a ventral production of the sternum 
is inhibited by the presence of a caudomesial boss on the coxa of this appendage; 
in D. (F.) carlsoni, however, the lateral lobes of the sternum are produced ven- 
tromesially much beyond the articulation between the coxa and the sternum. 

(5) The relative length and width of the carpus of the cheliped (Fig. Ip). In no 
member of the genus Cambarus is the length of the carpus so much greater than 
its width. 

(6) The cleft sternum anterior to the annulus ventralis (Fig. In). The narrow 
suture that lies between the sternal plates associated with the fourth pereiopods 
in D. (F.) carlsoni is not evident in any member of the genus Cambarus. 

(7) The articulation of the annulus ventralis. The hinge-like motion of the an- 
nulus ventralis is accomplished between the annulus and the sternum anterior to 
it rather than, as in Cambarus, in a transverse, weakly sclerotized area across 
the annulus proper. 

These differences seem sufficiently significant to conclude that this crayfish is 
not closely allied to members of any of the subgenera of Cambarus. Closer rel- 
atives seem to exist in the two members of the genus Distocambarus, for both 
Distocambarus (D.) devexus (Hobbs, 1981) and D. (D.) crockeri Hobbs and 
Carlson (1983) exhibit all of the features of carlsoni mentioned above. To be 
sure, there are marked differences in the form of the terminal elements of the 
first pleopod of the male, and in neither of the previously described species of 
Distocambarus is the central projection so elongate or dispased so distinctly 
caudally as it is in D. (F.) carlsoni. The similarities between the first pleopods 
of the latter (Fig. 1b, herein) and D. (D.) crockeri (see Hobbs and Carlson 1983: 
fig. 1b) are indeed striking, and a shortening of the shaft of the latter with a 
concomitant elongation of the central projection and reflection of the central 
projection would produce an appendage that would be little different from that 
of D. (F.) carlsoni. 

The assumption might be made that Distocambarus (F.) carlsoni links the 
genera Distocambarus and Cambarus. Assuming that the latter contains the more 
advanced species, if D. (F.) carlsoni links the two genera, and if one maintains 
the current concept of the origin of at least some Cambarus from an orconectoid 
ancestor (see Hobbs 1969), the conclusion might well be reached that the genus 
Cambarus as currently constituted has had a diphyletic origin. In my opinion, 
the Cambarus-like characteristics of D. (F.) carlsoni have been acquired inde- 
pendently. If such a conclusion is accepted then the genus Cambarus may be 
assumed to represent a group of species with a more recent common ancestry 
than any member has had with crayfishes assigned to other groups. But what 
recognition should be accorded the species described here? 

The fact that D. (F.) carlsoni shares the list of features just cited with the two 
species placed in Distocambarus by Hobbs and Carlson (loc. cit.) suggests that 


VOLUME 96, NUMBER 3 437 


assigning it to that genus is appropriate; however, there are several characteristics 
that set it apart from them: (1) The blade-like central projection on the first 
pleopod of the first form male. This feature, so strikingly resembling that typical 
of most members of the genus Cambarus, represents an apomorphic (lengthening 
of the terminal element) departure from the rather shorter and more distally 
oriented projection in the other two members of Distocambarus. 

(2) The restricted hinge motion of the annulus ventralis. Whereas in the two 
members of the subgenus Distocambarus the annulus is broadly hinged and swings 
through an arc of some 90 degrees, that of Fitzcambarus has an indistinct, narrow 
hinge anteriorly, and its arc of motion is no greater than 30 to 40 degrees. 

(3) The comparatively small postannular plate. This small wedge-like sclerite 
bears little resemblance to its conspicuous homologue in the subgenus Distocam- 
barus. In the two species belonging to the latter, this plate-like structure may 
extend anteriorly, covering the posterior part of the depressed annulus ventralis 
(in D. (D.) devexus), or project more ventrally when the annulus is depressed (in 
D. (D.) crockeri); in D. (F.) carlsoni it is not plate-like and does not extend so 
far ventrally as the annulus even when the latter is depressed. 

(4) The shorter, proportionately broader chela. The comparatively shorter, broad 
chela of D. (F.) carlsoni is cambaroid, resembling that of members of the sub- 
genus Depressicambarus, whereas that of the two members of the nominate 
subgenus is distinctly procambaroid, possessing an elongate palm and resembling 
the chelae of certain members of the subgenera Austrocambarus, Girardiella, and 
Ortmannicus. 

(5) The more extreme adaptations to a fossorial habitat. The rostrum of D. (F.) 
carlsoni 1s decidedly shorter than that in the two species included here in the 
nominate subgenus; the areola is conspicuously narrower; the abdomen is pro- 
portionately smaller, and the pereiopods are decidedly more stocky. 

As pointed out above, in my opinion, the affinities of this crayfish are clearly 
with the disjunct species pair that has been assigned to the genus Distocambarus. 
Because of the unique combination of characters demonstrated by his Procam- 
barus (Distocambarus) devexus, and believing it to be more closely allied to 
members of Procambarus than to those of other genera, Hobbs (1981) assigned 
it to that genus and erected for its reception the monotypic subgenus Distocam- 
barus signifying its remoteness from other species groups within the genus. With 
the discovery of a closely allied species, Distocambarus (D.) crockeri, Hobbs 
and Carlson (1983) believed that a better representation of the kinship of these 
with other crayfishes would be had if Distocambarus were elevated to generic 
rank. Taking cognizance of the unique features that these two crayfishes share, 
and suggesting the similarities and differences between them and D. (F.) carlsoni, 
I am proposing that the latter be assigned to the monotypic subgenus Fitzcam- 
barus within the genus Distocambarus. 

Ecological notes.—Distocambarus (F.) carlsoni is a primary burrower, and I 
suspect that it is rare that members of the species are found far from one of the 
mouths of its complex tunnel system. In the type-locality, it shares the low-lying 
swamp and seepage area with another primary burrower, Cambarus (Jugicam- 
barus) carolinus (Erichson, 1846:87), but the two seem to have partitioned the 
area on the basis of comparatively static and flowing groundwater. The occur- 
rence of the latter species in the swampy area may well be limited to sections 


438 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


where there is an active movement of the ground water, at least some of which 
emerges to the surface and flows toward the nearby creek; indeed, on 2 November 
1982, water was issuing from the mouths of the tunnels of two of the specimens 
obtained, and the other two individuals were taken from burrows out of which 
water trickled when they were opened. There on the eastern periphery of the 
swamp, the land rises more abruptly than on the more gentle slope where the 
burrows of Distocambarus (F.) carlsoni were found. One to three openings which 
are often surrounded by irregular piles of earthen pellets mark the domiciles of 
the latter crayfish. There seems to be no pattern to the configuration of the 
complex system of galleries, and no doubt any model that might have been in- 
stituted would have been modified through necessity by the mats of roots of trees 
among which the resulting tunnels are entwined. In most burrows, horizontal 
passageways radiate irregularly from one or two vertical tunnels that are suffi- 
ciently deep to penetrate the water table. Side passages that end blindly may 
extend in almost any direction from the principal galleries. As in burrows of most 
crayfishes that spend the greater part of their lives in the soil, those inhabited by 
females are more highly branched than those occupied by males. 

Both of the burrowing crayfishes found at this locality are infested with ento- 
cytherid ostracods. Cambarus (J.) carolinus harbors only one species, Harpa- 
gocythere georgiae Hobbs III (1965:163), whereas Distocambarus (F.) carlsoni 
is infested with two more wide ranging ones, Ankylocythere ancyla Crawford 
(1965:148) and Entocythere dorsorotunda Hoff (1944:332). The latter two ostra- 
cods have been found on a number of different crayfishes, but H. georgiae, which 
has been reported from a single locality, was obtained from a collection containing 
C. (J.) carolinus and C. (Depressicambarus) latimanus (LeConte, 1856). In the 
latter instance, whether or not both crayfishes bore this commensal is not known. 
Perhaps its host is limited to C. (J.) carolinus, the habitat of which is much more 
restricted than that of C. (D.) latimanus, a crayfish that tolerates a wide range 
of ecological conditions existing from the panhandle of Florida northward to 
northern North Carolina. 


Acknowledgments 


I am grateful to Paul H. Carlson and to Georgia B. Hobbs for their assistance 
in collecting the specimens on which this paper is based and to Denton W. Crock- 
er, Skidmore College, H. H. Hobbs III, Wittenberg University, and C. W. Hart, 
Jr., a Smithsonian colleague, for their criticisms of the manuscript. 


Literature Cited 


Crawford, E. A., Jr. 1965. Three new species of epizoic ostracods (Ostracoda, Entocytheridae) 
from North and South Carolina.—American Midland Naturalist 74(1): 148-154, 16 figures. 

Erichson, W. F. 1846. Uebersicht der Arten der Gattung Astacus.—Archiv ftir Naturgeschichte, 
Berlin 12(1):86—-103. 

Hobbs, Horton H., Jr. 1969. On the distribution and phylogeny of the crayfish genus Cambarus. 
In Perry C. Holt, Richard L. Hoffman, and C. Willard Hart, Jr., The distributional history of 
the biota of the southern Appalachians, Part I: Invertebrates — Virginia Polytechnic Institute, 
Research Division Monograph 1:93—178, 20 figures. 

——. 1981. The crayfishes of Georgia.—Smithsonian Contributions to Zoology 318:viii + 549 
pages, 262 figures. 


VOLUME 96, NUMBER 3 439 


, and Paul H. Carlson. 1983. Distocambarus (Decapoda: Cambaridae) elevated to generic rank, 
with an account of D. crockeri, new species, from South Carolina.—Proceedings of the 
Biological Society of Washington 96(3):420—428, 1 figure. 

Hobbs III, H. H. 1965. Two new genera and species of the ostracod family Entocytheridae, with 
a key to the genera.—Proceedings of the Biological Society of Washington 78(19): 159-164, 6 
figures. 

Hoff, C. Clayton. 1944. New American species of the ostracod genus Entocythere.—American 
Midland Naturalist 32(2):327—357, 33 figures. 

LeConte, John. 1856. Descriptions of new species of Astacus from Georgia.—Proceedings of the 
Academy of Natural Sciences of Philadelphia 7:400-402. 


Department of Invertebrate Zoology, Smithsonian Institution, Washington, D.C. 
20560. 


PROC. BIOL. SOC. WASH. 
96(3), 1983, pp. 440-451 


DISPOSITION OF THREE SPECIES OF ONISCOIDEA FROM 
WESTERN ATLANTIC SEASHORES (CRUSTACEA: 
ISOPODA: HALOPHILOSCIIDAE AND PHILOSCIIDAE) 


George A. Schultz 


Abstract.—Three species of terrestrial isopods from the seashore, Alloniscus 
compar Budde-Lund, Philoscia culebrae Moore, and Philoscia vittata Say, are 
discussed. The first two are placed in Vandeloscia Roman (family Halophilosci- 
idae), and P. vittata is placed in the new genus Sayoscia (family Philosciidae). 
Other related species are discussed briefly. Distribution and some general biology 
of the species are considered. 


Three species of terrestrial isopod crustaceans, Alloniscus compar Budde-Lund 
(1893), Philoscia culebrae Moore (1901) and Philoscia vittata Say (1818), were 
among those included by Van Name (1936) in his comprehensive work on isopods 
from the New World. The true identity of the three has never been adequately 
resolved (Lemos de Castro 1965; Schultz 1974; Ferrara 1974) so with additional 
morphological evidence they are discussed and reclassified here. Alloniscus com- 
par is considered to be too indefinitely known to be classified further, but records 
of specimens which have been called by Budde-Lund’s name are reconsidered 
and placed in Vandeloscia Roman (1977). Philoscia culebrae is also placed in 
Vandeloscia. The species Philoscia vittata is placed in a new genus. 

The two species of Vandeloscia are placed in the family Halophilosciidae 
(=Halophilosciinae Kesselyak, 1930, family Oniscidae as defined by Vandel 1962: 
474) because they have similar genital apophyses, among other characters, which 
are used to define members of the group. The subfamily has been considered to 
be a full family by Vandel (1973:27) followed by Ferrara (1974:207) and Ferrara 
and Taiti (1979:110). No formal or actual raising of it to the full family level 
appears in the literature. The nominate genus Halophiloscia was described in the 
tribe Halophilosciini of the family Oniscidae by Verhoeff (1908:521). Philoscia 
vittata, although in many ways similar to species of Vandeloscia, is retained in 
Philosciidae principally because it lacks a bilobed genital apophysis. 


Vandeloscia Roman, 1977 
Vandeloscia Roman, 1977:146.—Ferrara and Taiti, 1979:110. 


The genus was described ona species determined by Chelazzi and Ferrara (1978) 
and by Ferrara and Taiti (1979) to be a synonym of ‘‘Littorophiloscia’’ compar 
(Budde-Lund). The species ‘‘compar’’ has been placed in Philoscia, Alloniscus, 
Littorophiloscia, and *“‘Littorophiloscia’’ so that Roman’s description is the first 
time that “‘compar”’ as a species has been placed in a distinct genus. The inad- 
equately defined, erroneously used name Alloniscus compar Budde-Lund (1893) 


which has been used several times in the past remains an uncertain species. The 


VOLUME 96, NUMBER 3 44] 


species name of Strouhal (1966) has priority over that of Roman as will be shown 
below. Alloniscus compar of Vandel (1952) was adequately described and is iden- 
tical to Halophiloscia (Stenophiloscia) riedli Strouhal (1966) and to Vandeloscia 
orientalis Roman (1977). The taxonomic status of the species is discussed under 
V. riedli here. 

The genus was not defined in a separate section by Roman, who distinguished 
it from Halophiloscia Verhoeff by the setation of the palp of the maxilliped. 
Pleopod | of the male is used here to distinguish the genus and species rather - 
than the much less definitive arrangement of setae on the palp of the maxilliped. 
A brief diagnosis of the genus is given here based on the two species now in the 
genus. The diagnosis of Vandeloscia Roman is written with the criteria for defin- 
ing genera of Philosciidae, compiled by Vandel and listed by Taiti and Ferrara 
(1980:56), in mind. 

Diagnosis.—Pleon narrower than peraeon; edges on pleonal segments show 
slightly (dorsal view). Pigmentation, especially on edges of peraeonal segments 
and on peraeopods, conspicuously arranged in chromatophores. No glands, but 
tiny lateral nodes present on peraeonal segments. Cephalon with no frontal line; 
supra-antennal line well defined. Tip of genital apophysis strongly bilobed. En- 
dopod of male pleopod | with short, narrow apex strongly folded laterally. Pe- 
raeopod I sexually dimorphic. Dactylar organs present on peraeopods. 

Type-species.—Halophiloscia (Stenophiloscia) riedli Strouhal (1966), by mono- 
typy. 

Derivation and gender of name.—TYhe genus is named for the late French 
carcinologist, Dr. Albert Vandel. The gender is feminine. 

Other species.—Vandeloscia culebrae (Moore, 1901) (see below). 

Affinities. —The lobes on the tip of the genital apophysis are not as long in 
species of Vandeloscia as they are in species of Halophiloscia. 


Vandeloscia riedli (Strouhal, 1966) 
Fig. 1J-R 


Alloniscus compar Budde-Lund.—Vandel, 1952:112, figs. 30-33.—Arcangeli, 1958: 
242 —Cloudsley-Thompson, 1971:10.—Schultz, 1974:149 (as P. compar Budde- 
Lund, 1885).—Vandel, 1977:393.—Roman, 1977:135. 

Littorophiloscia compar (Budde-Lund).—Lemos de Castro, 1965:94, figs. 31- 
33.—Chelazzi and Ferrara, 1978:192.—Ferrara and Taiti, 1979:110. 

‘“‘Littorophiloscia’’? compar (Budde-Lund).—Ferrara, 1974:207, figs. 63-79. 

Halophiloscia (Stenophiloscia) riedii Strouhal, 1966:325, figs. 1-23. 

Philoscia (Setaphora) sp.—Roman, 1970:168. 

Vandeloscia orientalis Roman, 1977:146, figs. 20-23. 

Stenophiloscia riedli Strouhal.—Ferrara, 1974:212, fig. 82. 

nec Alloniscus compar Budde-Lund, 1893:124.—Dollfus, 1893:345.—Van Name, 
1936:218.—Arcangeli, 1960:47 (=incertae sedis). 


The species is the same as that described by Vandel (1952) and Lemos de 
Castro (1965) from the New World, and Ferrara (1974), Roman (1977) and Strou- 
hal (1966) from the Old World as comparison of their illustrations and those here 
will show. According to Arcangeli (1958:242) Vandel’s A. compar is certainly not 
conspecific with Budde-Lund’s A. compar. In fact there is no reason to believe 


442 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 1. A-—I, Vandeloscia culebrae: A, Male peraeopod I; B, Detail male peraeopod I; C, Male 
peraeopod VII; D, Female peraeopod I; E, Detail female peraeopod I; F, Female peraeopod VII; G— 
I, Male pleopods 1-3. J-R, Vandeloscia riedli: J-L, Male pleopods 1-3; M, Male peraeopod I; N, 
Detail male peraeopod I; O, Male peraeopod VII; P, Female peraeopod I; Q, Detail female peraeopod 
I; R, Female peraeopod VII. 


that it is the briefly described Alloniscus compar Budde-Lund (1893) which was 
not illustrated and which has as its type-locality ‘‘La Moka and vicinity of Ca- 
racas,’ Venezuela. The type-locality apparently is not a beach, nor near a beach, 
which is where all specimens like those described by Vandel (1952) and others 


VOLUME 96, NUMBER 3 443 


were collected. Until more specimens from the type-locality (the exact location 
of “‘La Moka”’ has yet to be determined) are available, A. compar Budde-Lund 
must be considered to be incertae sedis. 

From Budde-Lund’s brief Latin description without illustrations of A. compar, 
it can only be concluded that the species has eyes and three flagellar articles on 
antenna 2. The other characters of Budde-Lund’s species are so generally de- 
scribed that they might be present on any oniscoid. Vandel (1952), when he used 
Budde-Lund’s name for his specimens, made no mention of Budde-Lund’s brief 
description. Since the locations of the collections of other species included by 
Budde-Lund (1893) were not at the seashore, A. compar probably was not col- 
lected there or it would have been mentioned. The location where the specimens 
of Vandel were collected was ‘‘Orchilla, Iles-sous-le-Vent, a lest de Il’ Archipel 
de los Roques.” The location is a tiny islet and, although not directly stated by 
Vandel, the specimens most probably were taken on the seashore as were some 
other species included by Vandel. The only thing which the collection locations 
of the two species have in common (assuming that “‘vicinity of Caracas’’ is inland 
like the city) is that they are in Venezuela. 

Based on Vandel’s description, Arcangeli (1958:243) removed Alloniscus com- 
par of Vandel from Alloniscus Dana. Arcangeli stated that Vandel’s A. compar 
probably belonged to a different genus of philoscomorph, which he did not name; 
he continued, however, to keep A. compar in Alloniscus. Roman (1977:135) 
also included A. compar in her list of species of Alloniscus, never relating it to 
A. compar of Vandel. Ferrara and Taiti (1979:110) finally showed the similarities 
of Vandel’s and Roman’s specimens, but did not discuss any of the taxonomic 
implications. Vandeloscia riedli is not completely redescribed here because Van- 
del (1952), Ferrara (1974), Roman (1977), and Strouhal (1966) adequately illus- 
trated and described it. Details of male and female peraeopods I and VII and of 
male pleopods | and 2 are included here so the species can be easily distinguished 
from V. culebrae (Moore) with which it has been confused in the past. 

Diagnosis.—Tiny, but obscure, lateral nodes present on all peraeonal seg- 
ments. Endopod of pleopod 1 of male with large scalelike subapical process on 
laterally folded tip. Propodus of male peraeopod I with 2 large setae on inner 
margin. Exopod of pleopod 2 of male with 2 setae on posterolateral border. 

Description.—See Vandel (1952:112, figs. 30-33), Ferrara (1974:207, figs. 53- 
79), Roman (1977:146, figs. 20-23) and Strouhal (1966:325, figs. 1-23) for descrip- 
tions. Strouhal illustrated the small lateral nodes on peraeonal segment I of his 
specimens, and they were present but very obscure on the specimens examined 
here. More than one specimen had to be examined before they were discovered. 
Apparently lateral nodes were missed by Roman (1977). 

Measurements.—Males up to 5.9 mm long; females up to 6.0 mm long. 

Type-locality.—Isle of Abu Mengar, south of Ghardaga, in the Red Sea of 
Egypt. 

Distribution.—Gulf of Aquaba and Red Sea coast of Egypt; Suakin, Sudan; 
Sar Uanle and Gesira, Mogadishu, Somalia; Tulear, Madagascar; St. Helena Is- 
land, South Atlantic; Bay of Bengal, India; Yucatan, Mexico; San Pedro, Belize; 
Orchilla, Venezuela; Cabo Frio, Brazil. 

Type-specimens .—Strouhal (1966) makes no mention of the disposition of the 
type-specimens, but they are probably in the Natural History Museum in Vienna. 


444 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Material examined.—Yucatan, Mexico, and Belize, Central America. Speci- 
mens from both locations have been deposited in the National Museum of Natural 
History. 

Affinities. —See Affinities section under Vandeloscia culebrae. 


Vandeloscia culebrae (Moore, 1901) 
Fig. 1A-I 


Philoscia culebrae Moore, 1901:176, pl. 11, figs. 13—-17.—Richardson, 1905:604, 
fig. 660.—Boone, 1918:602.—Van Name, 1924:194, 195;—1936:168, fig. 86.— 
Vandel, 1968:114.—Schultz, 1974:149. 

Halophiloscia culebrae (Moore).—Vandel, 1945:242;—1949:8.—Arcangeli, 1948: 
482.—1958:242. 

Littorophiloscia compar culebrae (Moore).—Lemos de Castro, 1965:96, figs. 
36-39. 

Philoscia miamiensis Schultz, 1966:457, figs. 1-33. 

Alloniscus culebrae Moore, 1901.—Roman, 1977:137. 

nec Philoscia culebrae.—Pearse, 1915:534, fig. B. 


Moore (1901) gave a very general description of the two females which he 
collected on a beach on Culebra Island off Puerto Rico. The species generally is 
darker in color than V. riedli, but the chromatophores are still very conspicuous 
on the edges of the peraeonal segments and the peraeopods. The species is much 
like V. riedli, but differs from that species in the characters which are included 
in the diagnosis. 

Diagnosis.—Tiny lateral nodes present on peraeonal segments. Endopod of 
pleopod | of male with small scalelike subapical process on laterally folded tip. 
Propodus of male peraeopod I with 3 large setae on inner margin. Exopod of 
pleopod 2 of male with | seta on posterolateral border. 

Description.—See Schultz (1966) for overall view and Lemos de Castro (1965) 
for detail of male pleopods. Some differences or additions to the description 
include the fact that there are 9, not 21, ocelli, and tiny lateral nodes are present 
on the peraeopods. Even though the lateral nodes are tiny, they are more distinct 
than those of V. riedli. 

Measurements.—Males up to 5.0 mm long; females up to 6.1 mm long. 

Type-specimens.—Moore (1901) deposited his specimens (2 females) in the 
National Museum of Natural History. Very little is to be learned from them 
without dissection other than that they are similar when compared to fresh female 
specimens of the species from Cedar Key, Florida, and St. John, Virgin Islands. 

Type-locality.—Culebra Island (east of Puerto Rico). 

Distribution.—Culebra Island, Puerto Rico; Puerto Rico; St. John, Virgin Is- 
lands; east and west coasts of Florida. Abundant in maritime drift and other 
organic detritus on beaches. 

Material examined.—Cedar Key, west coast of Florida, and St. John, Virgin 
Islands. Specimens from each location have been deposited in the National Mu- 
seum of Natural History. 

Affinities.—It is difficult to tell females of the two species apart so no attempt 
is made here to distinguish them. Although the range of color of individuals in a 


VOLUME 96, NUMBER 3 445 


freshly preserved group of specimens of both species is great, the average density 
of color of fresh specimens of V. culebrae is greater. The two species can best 
be distinguished by comparing the size of the scalelike process on the tip of the 
endopod of male pleopod |. It is smaller and not always too distinct in V. cule- 
brae; whereas it is larger and divided into one large and one small process as 
shown here (Fig. 1J) for V. riedli. The propodus of male peraeopod I bears three 
large setae in V. culebrae and only two in V. riedli. Other consistent differences 
are present on other appendages and can be seen by comparing the illustrations 
given here and those of other workers. 

The differences present here which are used to distinguish the two species, in 
general, are valid if one also compares the two species as described from other 
locations or from the descriptions of other workers from the literature. Specimens 
of V. culebrae described by Schultz (1966, as Philoscia miamiensis) and Lemos 
de Castro (1965, as Littorophiloscia compar culebrae) have the same type of 
small scalelike process on the tip of the endopod of male pleopod 1. They also 
have the same corresponding configuration of setae on the inner margins of the 
propodus and carpus on male peraeopod I. 

'Specimens of V. riedli as presented in the illustrations of Vandel (1952, as 
Alloniscus compar), Lemos de Castro (1965, as Littorophiloscia compar compar), 
Strouhal (1966, as Halophiloscia (Stenophiloscia) riedli), Ferrara (1974, as **Lit- 
torophiloscia’’ compar) and Roman (1977, as Vandeloscia orientalis) have the 
same large scalelike process on the tip of the endopod of male pleopod 1. 
Correspondance between the configurations of other structures illustrated for the 
two species are also quite striking so that the two species can be said to be distinct 
both by using fresh specimens and by comparisons of illustrations and descriptions 
of the specimens in the literature. The differences are much more consistent than 
is to be expected at the subspecies level as was considered to be so by Lemos 
de Castro (1965). Vandeloscia culebrae is tropical in distribution, but it also 
ranges into subtropical Florida. Vandeloscia riedli is much more widespread on 
tropical beaches in the West Indies, Central America, and northern South Amer- 
ica and is common in parts of the Old World as well. 


Sayoscia, new genus 


The single species of the genus lives along the eastern shore of North America 
from Canada to Texas. The genus is placed in the broadly defined Philosciidae, 
not Halophilosciidae, because the type-species does not have the bilobed genital 
apophysis characteristic of the Halophilosciidae. When more species of the phi- 
loscomorph complex of species of oniscoids are adequately defined, better criteria 
for separating the Halophilosciidae from the Philosciidae will be distinguished. 
The species is darkly pigmented with distinct chromatophores on the peraeonal 
segments and on the peraeopods. The genital apophysis is without a bilobed tip, 
but has well developed grooves on the edges into which the very broad endopods 
of pleopod | fit. 

Diagnosis.—Peraeonal segments without glands, but with tiny lateral nodes 
(with seta about twice size of other body setae) on segments I-IV. Short apical 
extensions markedly bent laterally on tips of very broad endopod of male pleopod 
1. Peraeopods I, II, and VII similar in males and females. Dactylar organs present 


446 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 2. Sayoscia vittata: A, Dorsal view male 4.3 mm long; B, Frontal view cephalon; C, Tip of 
genital apophysis; D, Antenna 2; E, Antenna 1; F—J, Male pleopods 1-5; K, Pleotelson; L—N, Male 
peraeopods I, If and VII; O, Left mandible; P, Right mandible; Q, Maxilla 2 (endopod and exopod); 
R, Maxilliped; S, Maxilla 1; T, Uropod. 


on peraeopods. Mandibles with molar setae narrow at base with compound apex. 
Lacinia mobilis on each mandible. 

Type-species.—Philoscia vittata Say (1818). 

Derivation and gender of name.—The genus is named for Thomas Say, Amer- 


VOLUME 96, NUMBER 3 447 


ican entomologist and naturalist, who described the type-species. The gender is 
feminine. 

Affinities.—The single species of the new genus is similar to species of Van- 
deloscia Roman, but lacks the bilobed tip of the genital apophysis found in species 
of that genus. 


Sayoscia vittata (Say, 1818) 
Fig. 2A—F 


Philoscia vittata Say, 1818:429.—DeKay, 1844:50.—White, 1847:99.—Verrill and 
Smith, 1873:569.—Harger, 1879:157;—1880:306 (not pl. 1, fig. 1) —Budde-Lund, 
1885:209.—Underwood, 1886:361.—Richardson, 1900:305:—1901:565:—1905: 
605, figs. 662-663 (not fig. 661).—Paulmier, 1905:181 (not fig. 53).—Rathbun, 
1905:45, 4.—Fowler, 1912:233 (not pl. 66).—Sumner, Osborn, and Cole, 1913: 
661.—Pratt, 1951:442 (not fig. 611).—Kunkel, 1918:240 (not fig. 77)—Van Name, 
1936:115 (not fig. 52).—Schultz, 1974, fig. 3b, c, f;—1975:186;—1977: 154. 

Philoscia robusta Schultz, 1963:27, figs. 1-22;—1965:108;—1966:461. 

Philoscia (Philoscia) muscorum (Scopoli), var. sylvestris (Fabricius).—Blake, 
1931:351.—Procter, 1933:248. 

Philoscia muscorum (Scopoli).—Pratt, 1951:442. 

Littorophiloscia vittata (Say).—Alexenburg, 1962:23.—Bousfield, 1962:51.—Le- 
mos de Castro, 1965:90, figs. 21-30. 

partim Philoscia muscorum (Scopoli).—Harger, 1880:306 (pl. 1, fig. 1).—Schultz, 
1965: 107.—1974: 147. 


The species was redescribed by Schultz (1963) as Philoscia robusta. It was 
further discussed as P. vittata Say by Schultz (1974) who showed how it and P. 
muscorum (Scopoli), which shares part of its range, were confused by early 
workers. Indeed, the illustration of Harger (1880, pl. 1, fig. 1) of P. muscorum 
has been repeated by most other workers who encountered P. vittata on the east 
coast of the United States. The two species easily can be distinguished by com- 
paring male pleopods |. The illustrations of the species here are of specimens 
collected in south New Jersey not far from the type-locality as recorded by Van 
Name (1936). 

Diagnosis.—See generic diagnosis. 

Description.—Schultz (1966) completely described the species as Philoscia ro- 
busta and into that description the information in the generic diagnosis section 
can be incorporated. 

Measurements.—Males to 5.8 mm long; females to 6.0 mm long. 

Type-locality.—Great Egg Harbor, New Jersey (Van Name 1936), or according 
to Say (1818), “‘United States, common.”’ Say worked at the Academy of Natural 
Sciences, Philadelphia, and field trips frequently were taken to the New Jersey 
seashore. Say also toured much of the shore of the then United States and perhaps 
collected the species as far south as Georgia. 

Type-specimens.—Dry specimens were deposited in the Academy of Natural 
Science in Philadelphia, but they are no longer present (Lemos de Castro 1965: 
92). One dry specimen is present in the British Museum (Natural History) (White 
1847) apparently presented by Thomas Say himself. Types and other specimens 


448 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 3. Sayoscia vittuta: A-C, Female peraeopods I, II and VII; D-F, Female pleopods 1-3. 


of Philoscia robusta Schultz, a junior synonym, from near Beaufort, North Car- 
olina, are in the National Museum of Natural History and more specimens from 
New Jersey have been deposited there in connection with this redescription. 

Material examined.—The specimens examined here are from south Cape May 
County, New Jersey, on the edge of Grassy Sound. 

Distribution.—The species is present on the Atlantic coast of North America 
from the Bay of Fundy, Canada, to the mouth of the St. Marys River, Georgia. 
Specimens are present on the Gulf shore at least to the southern Texas coast. 
They are abundant in decayed organic vegetation, under the maritime drift in 
marine swamps, and in places regularly moistened by marine or estuarine waters. 

Disposition of some species of philoscomorphs from the New World which 
have been considered related to the three species considered above are those of 
the “‘Halophiloscia’’ group of Van Name (1936:167, 515). The species are Phi- 
loscia bermudensis, P. brasilensis, P. culebrae, P. culebroides, P. nomae, and 
P. richardsonae. Vandel (1945:242, 1949:8) gives maps of the distribution of the 
species of Halophiloscia Verhoeff including members of Van Name’s ‘‘Halophi- 
loscia’’ group. Vandel (1962:475) considered Littorophiloscia Hatch (1947) to be 
a junior synonym of Halophiloscia Verhoeff, but later (1968:113) he considered 
it to be valid with one species L. richardsonae (Holmes and Gay), the type- 
species. Lemos de Castro (1958a, 1965) included Al/oniscus compar Budde-Lund 
and Philoscia vittata Say and some of the above species of the “‘Halophiloscia”’ 
group as definitely or tentatively in Littorophiloscia Hatch. He based his conclu- 
sions on similarities of certain characters, especially general similarities of the 
shapes of the endopods of male pleopod |, to those of L. richardsonae. 

The disposition of all species considered in the ‘“‘Halophiloscia’’ group of Van 
Name (1936) and Littorophiloscia as expanded by Lemos de Castro (1965) is 
given below. Formal synonymies for species in the list and not included in syn- 
onymies above are to be found in the works mentioned after the valid name of 
the species. 


Philoscia bermudensis Dahl = Halophiloscia couchi (Kinahan); see Vandel (1962: 
477), and Schultz (1972:86). 


VOLUME 96, NUMBER 3 449 


Philoscia bonariensis Giambiagi de Calabrese = Halophiloscia couchi (Kina- 
han); see Reca (1972:407). 

Halophiloscia brasilenses Moreira = Benthana olfersi (Brandt); see Lemos de 
Castro (1958b:98), and Vandel (1962:475). 

Alloniscus compar Budde-Lund = incertae sedis; see text. 

Alloniscus compar Budde-Lund of Vandel (1952:112) = Vandeloscia riedli 
(Strouhal); see text. 

Philoscia culebrae Moore = Vandeloscia culebrae (Moore); see text. 

Philoscia culebroides Van Name = Nesophiloscia culebroides (Van Name); see 
Vandel (1968:113). 

Philoscia nomae Van Name = Nesephiloscia culebroides (Van Name); see Van- 
del (1968:113). 

Philoscia richardsonae Holmes and Gay = Littorophiloscia richardsonae (Holmes 
and Gay); see Lemos de Castro (1965:88), and Vandel (1968:113). 

Philoscia vittata Say = Sayoscia vittata (Say); see text. 


Acknowledgments 


I would like to thank Dr. Thomas E. Bowman of the National Museum of 
Natural History for reviewing the manuscript. I would also like to acknowledge 
the use of Separately Budgeted Research Funds from Jersey City State College 
for the project. 


Literature Cited 


Arcangeli, A. 1948. Il genere Halophiloscia Verhoeff nella regione mediterranea.—Archivo Zool- 
ogico Italiano 33:473-485. 

———. 1958. Le specie di Isopodi terrestri che furono erroneamente assegnate al genere Alloniscus 
Dana.—Memorias del Museo Civico di Storia Naturale di Verona 6:239-252. 

——. 1960. Revisione del genere Alloniscus Dana. I] sistema respiratorio speciale agli exopoditi 
dei pleopodi delle specie appartenenti allo stesso genere (Crostacei isopodi terrestri).—Bolle- 
tino Istituto e Museo di Zoologia dell’ Universita di Torino 6:17-79. 

Alexenburg, M. L. 1962. A key to the sowbugs of Long Island, New York.—Turtox News 40(1): 
22-23. 

Blake, C. H. 1931. Distribution of New England wood lice (on Isopoda Oniscoida, third paper).— 
Occasional Papers of the Boston Society of Natural History 5:349-355. 

Boone, P. L. 1918. Description of ten new isopods.—Proceedings of the United States National 
Museum 54(2253):591-604. 

Bousfield, E. L. 1962. Studies on the littoral marine arthropods from the Bay of Fundy region.— 
Bulletin of the National Museum of Canada 183:42-62. 

Budde-Lund, G. 1885. Crustacea Isopoda terrestria per familias et genera et species descripta.— 
Hauniae, pp. 1-319. 

———. 1893. Landisopoder fra Venezuela, indsamlede af Dr. Fr. Meinert.—Entomologiske Med- 
delelser, pp. 111-129. 

Chelazzi, G., and F. Ferrara. 1978. Researches on the coast of Somalia. The shore and the dune 
of Sar Uanle. 19. Zonation and activity of terrestrial isopods (Oniscoidea).—Monitore Zoolo- 
gico Italiano N.S. Supplemento XI, 8:189-219. 

Cloudsley-Thompson, J. L. 1971. The littoral terrestrial fauna of Suakin, Sudan.—Entomologist’s 
Monthly Magazine 107:10. 

DeKay, J. E. 1844. Zoology of New York, or the New York fauna. Part IV. Crustacea (Isopoda, 
pp. 42—54) Carroll and Cook: Albany. 

Dollfus, A. 1893. Voyage de M. E. Simon au Venezuela (Décembre 1887—Avril 1888).—Annales de 
la Société Entomologique de France 62:339-346. 


450 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Ferrara, F. 1974. Researches on the coast of Somalia. The shore and the dune of Sar Uanle. 3. 

Terrestrial isopods.—Monitore Zoologico Italiano N.S. Supplemento V, 15:191—220. 

, and S. Taiti. 1979. A check-list of terrestrial isopods from Africa (south of the Sahara).— 

Monitore Zoologico Italiano N.S. Supplemento VII, 1:1-114. 

Fowler, H. W. 1912. The Crustacea of New Jersey.—Annual Report of the New Jersey State 
Museum 1911:29-651. 

Harger, O. 1879. Notes on New England Isopoda.—Proceedings of the United States National 
Museum 2(79): 157-165. 

—. 1880. Report on the marine Isopoda of New England and adjacent waters.—Report of the 
United States Commissioner of Fish and Fisheries, for 1878. Part 6:297—462. 

Hatch, M. H. 1947. The Chelifera and Isopoda of Washington and adjacent regions.—University 
of Washington Publications in Biology 10:155—274. 

Kesselyak, A. 1930. Halophiloscinae n. subfam. ftir die Gattungen Halophiloscia und Stenophilos- 
cia.—Zoologischer Anzeiger 98:282—285. 

Kunkel, B. W. 1918. The Arthrostraca of Connecticut. Part II. Ilsopoda.—Connecticut State Geo- 
logical and Natural History Survey Bulletin 26: 185-261. 

Lemos de Castro, A. 1958a. On the systematic position of some American species of Philoscia 

Latreille (Isopoda, Oniscoidea).—American Museum Novitates 1908:1—10. 

1958b. Revisao do género Benthana Budde-Lund, 1908 (Isopoda, Oniscidae)—Arquivos 
do Museu Nacional Rio de Janeiro 46:85-118. 

—. 1965. On the systematics of the genus Littorophiloscia Hatch (Isopoda, Oniscidae).—Ar- 
quivos do Museu Nacional Rio de Janeiro 53:85—98. 

Moore, H. F. 1901. Report on Porto Rican Isopoda.—Bulletin Number 20, United States Fisheries 
Commission 2: 163-176. 

Paulmier, F.C. 1905. Higher Crustacea of New York City.—Bulletin New York State Museum 91: 
117-189. 

Pearse, A. S. 1915. An account of the Crustacea collected by the Walker Expedition to Santa 
Marta, Colombia.—Proceedings of the United States National Museum 49(2123):531—556. 

Pratt, H. S. 1951. A manual of the common invertebrate animals (exclusive of insects).—The 
Blakiston Co., Philadelphia, xviii + 854 pp. 

Procter, W. 1937. Biological survey of the Mount Desert region. Part V. Marine fauna.—Wister 
Institute of Anatomy and Biology, Philadelphia, 402 pp. 

Rathbun, M. J. 1905. Fauna of New England. 5. List of the Crustacea.—Occasional Papers of the 
Boston Society of Natural History 7:1—117. 

Reca, A. R. 1972. Oniscoideos Argentinos. II. Tres especies de isopodos terrestres de la costa 
maritima Bonaerense (Crustacea, Isopoda).—Physis 31:405—410. 

Richardson, H. 1900. Synopses of North-American invertebrates. VIII. The Isopoda.—Amenican 
Naturalist 34:207—230. 

—. 1901. Key to the isopods of the Atlantic coast of North America with descriptions of new 
and little known species.—Proceedings of the United States National Museum 23(1222):493— 
579. 

—. 1905. A monograph on the isopods of North America.—Bulletin of the United States Na- 
tional Museum 54:J—-LIII + 1-727. 

Roman, M.-L. 1970. Ecologie et répartition de certains groupes d’Isopodes dans les divers biotopes 
de la région de Tuléar (Sud-ouest de Madagascar).—Recueil des Travaux de la Station Marine 
d’Endoume, Marseille, Fascicule Hors Séries, Supplement 10: 163-208. 

———. 1977. Les oniscoides halophiles de Madagascar (Isopoda, Oniscoidea).—Beaufortia 26: 107— 
IS2, 

Say, T. 1818. An account of the Crustacea of the United States. By Thomas Say. Read September 
22, 1918 (Concluded).—Journal of the Academy of Natural Sciences of Philadelphia 1:423— 
444. 

Schultz, G. A. 1963. Philoscia robusta, a new species of terrestrial isopod crustacean from south- 
eastern United States.—Journal of the Elisha Mitchell Scientific Society 79:26—29. 

———. 1965. The reduction of Philoscia vittata Say, 1818, to a synonym of Philoscia muscorum 
(Scopoli, 1763).—Crustaceana 8:107—108. 

——.. 1966. Philoscia miamiensis n. sp., an isopod crustacean from Florida with ecological notes 
on the new species.—Transactions of the American Microscopical Society 85:457—462. 


VOLUME 96, NUMBER 3 451 


———. 1972. Ecology and systematics of terrestrial isopod crustaceans from Bermuda.—Crusta- 
ceana, Supplement 3:79-99. 

———. 1974. The status of the terrestrial isopod crustaceans Philoscia muscorum, P. vittata, P. 
robusta and P. miamiensis in the New World (Oniscoidea, Philosciidae).—Crustaceana 27: 
147-153. 

———. 1975. Terrestrial isopod crustaceans (Oniscoidea) from coastal sites in Georgia.—Bulletin 
of the Georgia Academy of Science 34: 185-194. 

———. 1977. Terrestrial isopod crustaceans (Oniscoidea) from St. Catherines Island, Georgia.— 
Georgia Journal of Science 35:151—158. 

Strouhal, H. 1966. Eine neue halophile Stenophiloscia aus dem Rotmeergebiete (Isop. terr.).— 
Annalen des Naturhistorischen Museum in Wien 69:323-333. 

Sumner, F. B., R. C. Osborn, and L. J. Cole. 1913. A biological survey of the waters of Woods 
Hole and vicinity. Part II, Section III. A catalogue of the marine fauna.—Bulletin of the United 
States Bureau of Fisheries 31:547—-794. 

Underwood, L. 1866. List of the described species of fresh water Crustacea from America, north 
of Mexico.—Bulletin of the Illinois State Laboratory of Natural History 2:323-326. 

Vandel, A. 1945. La répartition géographique des Oniscoidea (Crustacea Isopoda terrestres).— 
Bulletin Biologique de la France et de la Belgique 79:221-272. 

——. 1949. La fauna Nord-Atlantique.—Revue Francaise d’Entomologie 16: 1-11. 

———. 1952. Etude des Isopodes terrestres récoltes au Venezuela par le Dr. G. Marcuzzi.— 
Memorie dell Museo Civico di Storia Naturale di Verona 3:59-203. 

——. 1962. Isopodes terrestres (Deuxieme Partie).—Faune de France 66:417-931. 

. 1968. I. Isopodes terrestres. In Mission zoologique Belge aux iles Galapagos et en Ecuador 

(N. et J. Leleup, 1964—1965).—Musée Royal de |’ Afrique Centrale: Tervuren 1:37-168. 

——. 1977. La faune terrestre de l’ile de Sainte-Helene. Part 4. 1. Isopodes terrestres.—Annales 
Musée Royal de |’ Afrique Centrale, Série 8, Sciences Zoologiques 220:385—426. 

Van Name, W.G. 1924. Isopods from the Williams Galapagos Expedition.—Zoologica 5:181—210. 

1936. The American land and fresh-water isopod Crustacea.—Bulletin of The American 

Museum of Natural History 71:i-vii + 1-535. 

Verhoeff, K. W. 1908. Ueber Isopoden. 15. Isopoden-Aufsatz.—Archiv fiir Biontology 2:335—387. 

Vernll, A. E., and S. I. Smith. 1873. Report upon the invertebrate animals of Vineyard Sound and 
the adjacent waters, with an account of the physical characters of the region.—Report of the 
United States Commissioner of Fish and Fisheries for 1871-1872, part 8:295-778, pls. 1-38. 

White, A. 1847. List of the specimens of Crustacea in the collection of the British Museum.— 
Trustees of the British Museum: London, 143 pp. 


15 Smith St., Hampton, New Jersey 08827. 


PROC. BIOL. SOC. WASH. 
96(3), 1983, pp. 452-467 


AN UNUSUAL SPECIES COMPLEX IN THE GENUS 
EURYCOPE (CRUSTACEA: ISOPODA: ASELLOTA) 
FROM THE DEEP NORTH ATLANTIC OCEAN 


George D. F. Wilson 


Abstract.—A finely differentiated species flock or species complex in the asel- 
lote isopod genus Eurycope is described. The complex consists of four new species, 
plus three individuals that may belong to a fifth new species. The characters of 
these closely related species agree with the present definition of Eurycope, but 
several features of this complex make them distinct from their congenors. The 
most unusual of these traits is the very elongate stylet on the male pleopod II. 
The occurrence of this character in the Asellota and its phylogenetic significance 
are discussed. The elongate stylet is considered to be a derived trait that has 
evolved independently on numerous occasions in the Asellota Janiroidea. 


Introduction 


The genus Eurycope, recently revised by Wilson and Hessler (1981), contains 
numerous discrete species groups. Two such groups are the E. complanata com- 
plex and the E. inermis cluster (Wilson 1983). Here, another species complex is 
described within the genus which, while classifiable as Eurycope under the pres- 
ent system, is unique compared to its congenors. 

This group is designated the /ongiflagrata complex, after its most completely 
known species E. longiflagrata n. sp. The species of the complex all fit the present 
diagnosis of Eurycope, but are distinct in several traits from the other species in 
the genus. All males have a very distinctive external sexual apparatus: extremely 
long and whiplike stylets of the second pleopods that lie external to the pleopodal 
cavity, sometimes in grooves on the pleotelson dorsal surface. The body is gen- 
erally longer and narrower than in typical species of Eurycope. The medial length 
of pereonite 4 is less than that of pereonites 1-3, in contrast to the typical situation 
of 4 anterior pereonites of similar length. The body depth is also unusual: deepest 
at pereonite | instead of at pereonite 5 or 6 as generally found within Eurycope. 

The complex of species described here illustrates problems common in the 
study of deep-sea asellotes. 

1. Even though the genus Eurycope has been revised, with a number of species 
removed to other genera, many species in the genus are poorly known, creating 
uncertainty about additional divisions of the genus. The characters of the longi- 
flagrata complex fall within the definition of the genus, but its possession of 
unusual characters opens the possibility of the creation of a new genus, or the 
division of Eurycope into subgenera. Lack of knowledge prevents making a clear 
decision, so this complex is retained in Eurycope for the present. 

2. Separation of the species of the longiflagrata complex is often difficult, 
because they are rare in deep-sea samples. Consequently, there are few individ- 
uals for a study of character variation, making range overlaps, where they occur, 
difficult to interpret. In addition, females are difficult to classify because the 


VOLUME 96, NUMBER 3 453 


species are very similar and the most definitive characters are found in the males. 
Therefore, one species is described here on a provisional basis, with the hope 
that additional material will appear which will clarify some of the problems re- 
maining. Three specimens are described without any specific designation, al- 
though they may belong to a fifth new species. 


Materials and Methods 


The specimens on which this paper is based came from two sources. A long- 
standing deep-sea benthic ecology program at the Woods Hole Oceanographic 
Institution (WHOI samples), directed by H. L. Sanders, J. F. Grassle, and pre- 
viously R. R. Hessler, has provided isopods collected on sampling transects on 
most of the major basins of the Atlantic Ocean. The Centre National de Tri 
d’Océanographie Biologique, directed by M. Segonzac is the second important 
source of specimens. This French national sorting center has kindly sent isopods 
that were collected on three major oceanographic expeditions: Intercalibration 
(INCAL samples), led by M. Sibuet; Abyplaine (ABYPL samples), led by C. 
Monniot; and Demeraby (DEMERABY samples), led by M. Sibuet and C. Mon- 
niot. 

Types have been deposited at the United States National Museum of Natural 
History (cited as USNM). Remaining paratype and non-type specimens are re- 
tained in the research collection of Robert R. Hessler, Scripps Institution of 
Oceanography (RRH). 

The methods and terminology used in this paper are those developed by Wilson 
and Hessler (1980) and Wilson (1983). Measurement characters are generally 
stated as ratios to factor out allometric variation; these values are given to a 
precision of two significant figures. Because the sample sizes are very small, no 
statistical accuracy is implied for these values, although ranges are given where 
measurements for more than one individual were available. Descriptions are based 
on adult type-specimens. 

In the figures, mouthparts (except the mandible), pleopods, and uropods are 
shown in ventral view. Arrows indicate either enlarged sections of a limb, or 
structures of special interest which are labelled by a letter. In some cases, all 
setae are not shown but are indicated by u-shaped or circular marks at their 
insertions. 


Systematics 


Eurycope Sars, 1864 
The Eurycope longiflagrata complex 


Type species.—Eurycope longiflagrata n. sp. 

Diagnosis.—Eurycope with elongate body, body length approximately 3 times 
body width, body deepest at pereonite 1. Rostrum of cephalon short, narrow, 
anteriorly rounded, often overhanging frons, lacking cephalic keels but with small 
stout setae. Cephalic frons sloping anteriorly in front of rostrum; frons-clypeal 
ridge distinct, rounded, not shaped like an inverted V, often projecting anteriorly 
above clypeal insertion. Posterolateral corner of mandible inserting on cephalon 
into distinct notch (Fig. 1C, D-N). Pereonite 4 dorsomedial length reduced com- 


454 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


pared to other pereonites. Male sexual morphology modified: pleopod I distally 
u- or v-shaped with lateral lobes extending posteriorly and medial lobes reduced 
or absent; pleopod II stylet extremely elongate, about 3-4 times protopod length 
in adult; pleotelson dorsal surface often with groove or incision beginning on 
posterolateral margin behind uropod insertions and extending anteriorly (Fig. 1F— 
G). Uropodal protopod not broadened medially; rami longer than protopod width. 

Description.—Cephalon posterior to antennulae shorter than and almost as 
deep as pereonite 1. Lateral spine blunt and broad. Rostrum separated from 
cephalic dorsal surface by transverse depression; male rostrum often longer and 
narrower than that of female. 

Pereonite 7 and pleon lateral margins flattened, with cuticular ridges. Pereonite 
1 with greatest medial length of ambulosomites, lengths decrease to pereonite 4. 
Dorsal surfaces with scattered fine setae. Pereonite 7 longest natasomite, per- 
eonite 6 shortest. Pereonite 7 ventrolateral area anterior to coxae of pereopod 
VII lacking bulla, with single large seta anterior to coxa; posterior margin of 
pereon with dense row of plumose setae. 

Pleotelson lateral margin rounded, with anterior corners curving medially. 

Antennulae approximately one-third body length; more robust, longer, and with 
more flagellar articles in male than in female. Article | medial lobe short, not 
protruding, shorter than article 2; lateral plate rounded, not angular or protruding, 
with unusually large broom seta on lateral margin. 

Mandible typical, with palp shorter than mandibular body length and condyle 
shorter than molar process. 

Maxilliped with broad palp, palp article 4 medial lobe much shorter than article 
5. Epipod longer than wide, slightly shorter than basis, distally pointed; lateral 
projection small, angular. 

Pereopod I-II bases broader than in pereopods III-IV. Coxal plate of pereopod 
IV short, rounded, not projecting anteriorly as in more anterior pereonites. 

Pereopod V—VII bases slightly different in length: basis VI longest, basis V 
shortest. Carpus V—VII widest at less than quarter distance from distal edge. 

Pleopod II of female deep, with broad keel; posterior margin rounded or flat- 
tened. Apex variably expressed. Lateral margins with plumose (not hemiplumose) 
setae. 

Pleopod II of male with subterminal exopod having brush of long thin setae on 
posterolateral rounded edge. 

Uropodal protopod with long unequally bifid setae on ventral and distal margin. 
Endopod and exopod with distal rosette of robust unequally bifid setae. 

Remarks.—Members of the /ongiflagrata complex are most easily identified by 
their specialized male pleopod morphology, the somewhat narrowed body, the 
reduced dorsomedial length of pereonite 4 and by the sloping and protruding frons 
of the cephalon. The condition of the uropod is very similar to that seen in E. 
cornuta. 

The elongate stylet on the endopod of the male second pleopod is the most 
unusual feature of the /ongiflagrata complex. This type of stylet is not found in 
other Eurycopidae, but it makes scattered appearances in various forms on the 
species level in other Janiroidean genera. The advanced deep-sea genera in which 
this character is found are Munnopsis, Ischnomesus, Haploniscus, Acanthomun- 
na, and Dendromunna. Illustrations of this condition may be found in Sars (1899), 


VOLUME 96, NUMBER 3 455 


Wolff (1962), and Menzies (1962). An elongate stylet also occurs frequently in 
the more primitive janiroideans: Acanthaspidia, Ianthopsis, Ectias, Neojaera, 
and Caecianiropsis (see Hansen 1895; Kensley 1977; Nordenstam 1933; Schultz 
1976; Sivertsen and Holthuis 1980; Menzies and Pettit 1956). One could speculate 
that this type of stylet is a primitive character in the Janiroidea because of its 
widespread occurrence, and because of its appearance in the primitive groups. 
On the other hand, the stylet is quite variable in the genera in which it is elon- 
gated. For example, in Munnopsis one sees a range from an elongate stylet (M. 
typica) to a highly reduced stylet (M. abyssalis). In the very primitive genus 
Notasellus, which might be considered to be an outgroup to the more advanced 
Janiroidea by virtue of its non-opercular second pleopods, the stylet is not elon- 
gate. Therefore, the elongate stylet must be an apomorphy that appears conver- 
gently in many diverse janiroidean groups, comparable to the loss of the seventh 
pereopod in adults (Wilson 1976). The /ongiflagrata complex is unusual in that 
this character has become established in all of its species. 


Eurycope longiflagrata, new species 
Figs. 1, 2 


Material examined.—Holotype preparatory female, 3.8 mm, USNM 195071. 
Paratypes: copulatory male, 3.2 mm, USNM 195072; 8 individuals, RRH. Type- 
locality WHOI 131, 36°28.9'N, 67°58.2'W, 2178 m. Other material: Juvenile fe- 
male, WHOI GH4, 39°29'N, 70°34'’W, 2469 m, RRH. 

Distribution.—Continental slope off New England, USA, 2178-2469 m. 

Etymology.—Longiflagrata 1s a Latin compound adjective meaning ‘‘with long 
whip.’ 

Diagnosis.—Adult body length approximately 3-4 mm. Rostrum not over- 
hanging frons; medial part of frons ridge low, sloping smoothly into clypeal ar- 
ticulation. Pleotelson of male with dorsal groove extending no further than an- 
terior articulation of uropod. Male pleopod I with tapering lateral lobes, medial 
region smoothly v-shaped, medial lobes not expressed; distance from dorsal ori- 
fice to distal tip 0.27 pleopod I length. Female pleopod II with definite apex 
approximately quarter pleopod length from distal tip. Uropodal endopod length 
1.5 or greater protopod width. 

Description.—Body characters (Fig. 1!A—B, E-F): Length 2.7-2.9 width. In 
holotype, pereonite 1 depth 0.3 length. In female, pleotelson round posteriorly. 
In male, pleotelson distal tip folds down abruptly, somewhat flattened in dorsal 
view. 

Cephalic rostrum (Fig. 1C—D, G): Anterior tip with 2 small but stout setae, 
rostrum length-cephalic width ratio 0.15 in female, 0.19 in males (2 measured). 

Antennula (Figs. 1G, 2A): Length 0.3-0.4 body length, longest and most robust 
in males. Flagellum (distal to article 4) with 7-11 annuli and 2 distal aesthetascs 
in females, and 14-25 annuli, each supplied with single aesthetasc, in males. 
Article 2 length 0.5 medial length of article | in females, 0.7 in males. Article 3 
length 1.3—1.6 article 2 length. 

Mandible (Fig. 2B—D): Left incisor process with 3 subequal teeth, right incisor 
process with 4 irregularly spaced teeth. Lacinia mobilis with 6 short teeth. Spine 
row with 4-6 members. Molar process distal surface oval, with only low bumps 


456 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 1. Eurycope longiflagrata, types: A-B, I-J, 3.8 mm holotype preparatory female; C, H, 3.6 
mm copulatory male; D, brooding female fragment; E—F, 3.2 mm copulatory male; K—L, 3.0 mm 
preparatory female. A-B, Holotype, lateral and dorsal views, scale bar 1.0 mm; C—D, Cephalon, 
oblique and lateral views, antenna and antennula removed, n = notch at posterior part of mandibular 
insertion; E, Copulatory male, dorsal view, scale bar 1.0 mm; F, Pleotelson and pereonite 7, lateral 
view, g = groove in dorsal surface; G, Male cephalon, dorsal view; H, Male pleopod II, ventral view; 
I, Lateral margin of natasomites, pereopod VII intact, in plan view; J, Female pleotelson, ventral 
view; K, Female pleopod II, lateral view; L, Uropod, ventral view. 


VOLUME 96, NUMBER 3 457 


Fig. 2. Eurycope longiflagrata, paratypes: A, D, H-I, 3.6 mm copulatory male; B—C, E-G, J-L, 
3.0 mm preparatory female. A, Antennula; B—D, Mandible; B, Dorsal view; C, Medial view; D, Male 
palp; E, Paragnaths; F, Maxillula; G, Maxilla; H, Maxilliped, with enlargements of endite distal tip 
and seta on medial margin of palp article 3; I, Male pleopod I, spermatophore protruding from dorsal 
orifice, with enlargement of lateral lobe; J-L, Pleopods III-V. 


and 4—6 setae on posterior edge. Condyle length 0.19 mandibular body length. 
Palp second article length 0.51 mandibular body length. 

Maxilliped (Fig. 2H): Endite with 5 small and | large fan setae distally, and 5 
coupling hooks medially; lateral part of distal margin with one bicuspid projection. 
Palp articles 2 and 3 medial to lateral length ratios 0.67, 3.4 respectively. Epipod 
length 0.91 basis length, length 2.2 width. 

Pleopod I of male (Fig. 21): Length 4.5 width at dorsal orifice. Ventral surface 
with 2 plumose setae, and 2 rows of fine setae near each distal tip. Lateral lobe 
tip somewhat truncate, sloping anteromedially, with fine setae; dorsal part of 
lateral lobe not curling into ventral view. 

Pleopod II of male (Fig. 1H): Protopod length 1.8 width; lateral margin with 4 
plumose setae. Endopod length including stylet 5.1 protopod length. 

Pleopod II of female (Fig. 1J, K): Depth 0.31 length. Lateral margin with 2 
plumose setae. 


458 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Uropod (Fig. 1L): Endopod width 0.25 length. Exopod length 0.75 endopod 
length. Protopod ventral surface with circa 3 setae. 

Remarks.—Males of Eurycope longiflagrata are most easily recognised by their 
short dorsal grooves on the pleotelson and smoothly tapering lateral lobes on 
pleopod I. As typical in all longiflagrata complex species, the females of E. 
longiflagrata are difficult to separate from the other species. The frons-rostral 
characters, and the female pleopod II shape are most useful to identify female 
specimens that are in good condition. At present this species is known only from 
the continental slope off New England at 2178—2469 meters. 


Eurycope friesae, new species 
Fig. 3 


Material examined.—Holotype copulatory male, 4.2 mm, USNM 195073. Para- 
type brooding female, 4.7 mm, RRH. Type-locality WHOI 156, 0°46.0-46.5’S, 
29°28 .0-24.0'W, 3459 meters. Other material: Preparatory female, DEMERABY 
29, 8°09.2-10.2'N, 49°04.4—04.8’W, 4430 m, RRH. 

Distribution.—Equatorial Atlantic, 3459-4430 m. 

Etymology.—In honor of my wife, M. Katherine Fries- Wilson, whose constant 
encouragement has made important contributions to this work and to my research 
in general. 

Diagnosis.—Adult body length approximately 4-5 mm. Rostrum not over- 
hanging frons; medial part of frons ridge low, sloping smoothly into clypeal ar- 
ticulation. Pleotelson of adult male with elongate dorsal groove, extending well 
anterior to insertion of uropod; groove length greater than two thirds pleotelson 
length. Male pleopod I with narrow elongate lateral lobes, medial region u-shaped, 
medial lobes expressed by presence of 3-4 simple setae; distance from dorsal 
orifice to distal tip 0.34 total pleopod length. Female pleopod II with rounded 
keel and no distinct apex. Uropodal endopod length |.5 protopod width. 

Description (in ranges, holotype male value stated first, female second).—Body 
characters (Fig. 3A—C): Length 3.0—3.1 width. In holotype, pereonite | depth 0.23 
body length. Pleotelson posterior to uropods obtusely angular in dorsal view. 

Rostrum (Fig. 3E): Anterior margin with 4 stout and 2 fine setae. Length 0.19- 
0.16 cephalon width. 

Antennula (Fig. 3E—F): Length—body length ratio 0.34—0.29. Flagellum with 20— 
11 annuli. Article 2 length 0.70—0.62 medial length of article 1. Article 3 length 
1.8—1.4 article 2 length. 

Left mandible (Fig. 3G—I): Spine row with 8 members. Molar process triturating 
surface with 11 setae and numerous distinct teeth on posterior margin. Condyle 
length 0.17 mandibular body length. Palp second article length 0.51 mandibular 
body length. 

Maxilliped (Fig. 3J): Endite distal tip with tiny pointed denticles laterally. Epi- 
pod length 0.95 basis length; length 2.2 width. 


_" 


Fig. 3. Eurycope friesae, types: A-B, K-M, 4.2 mm holotype copulatory male; C-J; 4.7 mm 
paratype brooding female; A-B, Holotype, dorsal and lateral views, scale bar 1.0 mm; C, Female 
paratype, dorsal view, scale bar 1.0 mm; D, Pleotelson, lateral view; E, Cephalon, dorsal view; F, 


VOLUME 96, NUMBER 3 459 


Antennula, lateral view; G—I, Left mandible; G, Incisor process and spine row, ventral view; H, 
Incisor process and molar process, anterior view; I, Molar process, medial view, with enlargement 
of sensory pore; J, Maxilliped with enlargement of endite distal tip; K-L, Male pleopod I and en- 
largement of lateral lobe, respectively; M, Uropod. 


460 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Pleopod I of male (Fig. 3K—L): Length 4.2 width at dorsal orifice. Ventral 
surface with 2 rows of 5—6 plumose setae each. Distal tip of lateral lobes sloping 
anteromedially with fine setae in row on lateral margin and tiny denticles on 
medial margin; dorsal part of lateral lobe curling into ventral view. 

Uropod (Fig. 3M): Endopod width 0.24 length. Exopod length 0.80 endopod 
length. Protopod ventral surface with circa 9 setae; dorsal surface of male with 
medial row of 8 setae. 

Remarks.—The male of Eurycope friesae is the most strikingly modified of the 
longiflagrata complex: the stylet groove in the pleotelson is very long and dis- 
tinct. The females, which are similar to those of E. /ongiflagrata, can be recog- 
nised by the rounded keel of the second pleopod. The shape of the frons and 
rostrum separates E. friesae from the other known species of the complex. This 
species has been collected from the southern part of the Demerara Abyssal Plain 
and from the equatorial Atlantic deep sea. 


Eurycope hessleri, new species 
Fig. 4 


Material examined.—Holotype copulatory male, USNM 195074. Paratypes: 
Brooding female, 3.5 mm, USNM 195075; 8 damaged or fragmentary individuals, 
RRH. Type-locality WHOI 334, 40°42.6-44.0’N, 46°13.8-14.6’W, 4400 meters. 

Distribution.—Eastern Sohm Abyssal Plain in the North Atlantic Ocean, 
4400 m. 

Etymology.—In honor of Dr. Robert R. Hessler, who has made numerous 
important contributions to deep-sea systematics, and who has been a valued 
mentor, colleague, and friend to me. 

Diagnosis.—Adult body length approximately 3-4 mm. Rostrum overhanging 
frons; medial part of frons ridge rounded in lateral view, protruding over clypeal 
insertion. Pleotelson of male without dorsal stylet groove. Male pleopod I with 
short dorsally curling lateral lobes, medial region sinusoidally v-shaped in ventral 
view with numerous fine setae anteriorly; distance from dorsal orifice to distal 
tip 0.23 pleopod I length. Female pleopod II with rounded keel, definite apex, 
and flattened posterior surface; distance from apex to distal tip 0.3 total pleopod 
length. Rami of uropod comparatively short, endopod length 1.2 protopod width. 

Description.—Body characters (Fig. 4A—B, D): Length 3.0 width. Pleotelson 
distal tip rounded in dorsal view; anterior corners not curving medially. 

Rostrum (Fig. 4E): Anterior tip with 2 small stout setae and several simple 
setae. Length 0.15 cephalic width. 

Antennula (Fig. 4F—G): Proximal article lengths not sexually dimorphic (ratios 
apply to both male and female types). Article 2 length 0.6 medial length of article 
1. Article 3 length 1.25 article 2 length. 

Left mandible (Fig. 4H—K): Spine row with 6 members. Distal surface of molar 
process with numerous elongate teeth and 7 setae on posterior margin. Condyle 
length 0.17 mandibular body length. Palp article 2 length 0.53 mandibular body 
length. 

Maxilliped (Fig. 4M): Endite with 4 coupling hooks; lateral part of distal margin 
with 2 triangular projections. Palp article 2 medial margin with 3 setae, medial 
length 0.61 lateral length. Epipod length 0.96 endite length. 


VOLUME 96, NUMBER 3 461 


Fig. 4. Eurycope hessleri, types: A-C, F, 3.2 mm holotype copulatory male; D-E, G, Q, 3.5 mm 
brooding female; N—P, Copulatory male pleotelson fragments; H—M, R, 2.9 mm copulatory male. 
A, Male, dorsal view, scale bar 1.0 mm; B, Male pleotelson, lateral view; C, Cephalon, lateral 
view; D, Dorsal view of female; E, Cephalon, lateral view, antennula and antenna removed; F-G, 
Male and female antennulae, respectively; H-K, Left mandible; H, Dorsal view; I, Incisor process 
and lacinia mobilis, posterior view; J, Spine row, dorsal view; K, Molar process, anterior view; L, 
Right mandible, incisor process, posterior view; M, Maxilliped with enlargement of endite distal tip; 
N-O, Male pleopod I; N, Ventral and lateral views; O, Distal tip with enlargement of lateral lobe; P, 
Male pleopod II; Q, Female pleopod II, lateral view; R, Uropod. 


462 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Pleopod I of male (Fig. 4N): Length 4.1 width at dorsal orifice. Ventral surface 
with 2 rows of plumose setae, 4-5 setae per row in adults, 0-2 in juveniles; 2 
rows of simple setae on posterior third of ventral surface. Distal groups of simple 
setae on lateral base of lateral lobes, on tip of lateral lobes, and on medial region. 
Lateral lobes distally rounded with dorsal surface strongly curving into ventral 
view. 

Pleopod II of male (Fig. 4P): Protopod length |.8—2.0 width; lateral margin with 
3 plumose setae. Endopod (including stylet) length 3.8 protopod length. 

Pleopod II of female (Fig. 4Q): Depth 0.27 length. Lateral margin with 2-4 
plumose setae. 

Uropod (Fig. 4R): Endopod width 0.37 length. Exopod length 0.89 endopod 
length. Protopod ventral surface with circa 7 setae; medial edge of dorsal surface 
with row of 4 setae in male only. 

Remarks.—Eurycope hessleri has a small but distinct rostral overhang, and 
rather short and stocky rami of the uropods; these characters are sufficient to 
distinguish E. hessleri from E. longiflagrata and E. friesae. The males are distinct 
in lacking a dorsal groove in the pleotelson, and in having short, dorsally curled 
lateral lobes on the pleopod I distal tips. E. hessleri is known only from the type- 
locality in the eastern part of Sohm Abyssal Plain. 


Eurycope errabunda, new species 
Fig. 5 


Material examined.—Holotype preparatory female, 6.2 mm, USNM 195076. 
Paratypes: head fragment, pleotelson fragment, WHOI 330, RRH. Type locality: 
WHOIT 330, 50°43.5—43.4’N, 17°51.7—52.9'W, depth 4632 meters. Other Material: 
possibly male head fragment and juvenile male with choniostomatid copepod 
parasite, 4.7 mm, WHOI 287, 13°16.0-15.8’N, 54°52.2—-53.1’W, 4980-4934 m. Three 
individuals: manca 3, instar 4, and fragmentary female, WHOI 334 (see E. hessleri 
for data). Cephalon and anterior pereon fragment, ABYPLAINE CP19, 43°00’N, 
14°02.9-04.3'W, 5280 m. 

Distribution.—Widely scattered localities in the abyssal regions of the North 
Atlantic Ocean, 4632-5280 m. 

Etymology.—Errabunda is a Latin adjective meaning ‘“wandering.”’ 

Diagnosis (adult male characters not known).—Adults large, body length ex- 
ceeding 6 mm. Cephalic rostrum overhanging frons; frons ridge medially over- 
hanging clypeal articulation. Female pleopod II with distinct apex approximately 
quarter pleopod length from distal tip. Uropodal endopod length 1.8 protopod 
width (1.6—1.7 in specimens from non-type localities). 

Description of type-material.—Body characters (Fig. 5A—B): Body narrow, 
length 3.2 width, depth at first pereonite 0.22 body length. Pleotelson rounded 
posteriorly. 

Rostrum (Fig. 5C—D): Length 0.16 cephalic width. Anterior tip with 2 stout 
setae and several fine setae. 

Antennula (Fig. SF): Article 2 length 0.7 medial length of article 1. Article 3 
length 1.7 article 2 length. 

Left mandible: Spine row with 12 members. Molar process with 14 distal setae. 
Condyle length 0.18 mandibular body length. Palp article 2 length 0.49 mandibular 
body length. 


VOLUME 96, NUMBER 3 463 


oh) 


es 


Fig. 5. Eurycope errabunda: A-B, E-F, H, 6.2 mm holotype preparatory female; C—D, Paratype 
female cephalon fragment; G, K, Adult male head fragment, WHOI 287; I, 4.7 mm juvenile male, 
WHOI 287; J, Instar 4 female, WHOI 334. A—B, Holotype, dorsal and lateral views; C-D, Cephalon, 
oblique and lateral views, antennula and antenna removed; E, Pereonites 2—7, oblique ventral view, 
pereopod bases in plan view; F—G, Female and male antennulae, respectively; H-J, Uropods, com- 
parison of individuals from WHOI 330, WHOI 287, and WHOI 334, respectively; K, Maxilliped. 


Uropod (Fig. 5H): Endopod width 0.21 length. Exopod length 0.77 endopod 
length. Protopod ventral surface with circa 9 setae. 

Description and variation in specimens from other localities. —WHOI 334, in- 
star 4 juvenile female (Fig. 5J): Uropodal endopod 1.6 protopod width. Endopod 
width 0.23 length. Exopod length 0.83 endopod length. Protopod ventral surface 
with 3 setae. 

WHOI 287, parasitized juvenile male (Fig. 51): Uropodal endopod 1|.7 protopod 


464 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 6. Eurycope sp., longiflagrata complex, 4.7 mm preparatory female, ABYPL DS11: A, ceph- 
alon, dorsal view; B, cephalon, lateral view, antennula and antenna removed; C, Pleopod II, lateral 
view, with enlargement of dorsal margin; D, Antennula; E, Left mandible; F, Maxilliped with en- 
largement of endite distal tip; G, Uropod. 


width. Endopod width 0.22 length. Exopod length 0.80 endopod length. Protopod 
ventral surface with 6 setae; dorsomedial surface with | seta. 

WHOI 287, head fragment, possibly male (Fig. 5G): Antennular article 2 length 
0.81 medial length of article 1; article 3 length 1.92 article 2 length; aesthetascs 
on proximal annuli of flagellum (evidence for male classification). Maxillipedal 
endite with 7 small and one large fan setae distally, and 5 coupling hooks medially; 
lateral part of distal margin with quadrate projection. Maxillipedal epipod length 
0.93 basis length; length 2.0 width. 

Remarks.—The individuals classified as Eurycope errabunda are recognizably 
different from the other species in the /ongiflagrata complex: all are large, and 
have a distinctive configuration of the frons and rostrum. Because the adult male 
pleopods are unknown it is uncertain whether FE. errabunda represents one or 
several species. The variation observed in the uropods and in the antennulae of 
specimens from the different localities could be explained by ontogenetic changes 
or sexual dimorphism (Wilson 1981), but differences characteristic of geograph- 
ically-separated populations cannot be ruled out. Therefore, E. errabunda is pro- 
visionally considered to be a single species pending the collection of additional 
material and characterization of the male pleopods. 


VOLUME 96, NUMBER 3 465 


Fig. 7. Eurycope sp., longiflagrata complex. 3.5 mm copulatory male, INCAL 45: A, Antennula; 
B—D, Left mandible; B, Dorsal view; C, Incisor process and lacinia mobilis, posterior view; D, Molar 
process anterior view; E, Pleopod I, distal tip; F, Pleopod II; G, Uropod. Copulatory male fragment, 
estimated body length 3.7 mm, WHOI 330: H, Pleopod I with enlargement of distal tip; I, Pleotelson, 
lateral view. 


Eurycope sp. (spp.?) 
Figs. 6—7 


Three individuals in the available material of the /ongiflagrata complex were 
not assignable to any of the species described above. The specimens occur at 
three different localities and are damaged or fragmented. At least one new species 
is represented in this material, but because of the condition and ambiguities in 
character states, a new species is not proposed here. The individuals are de- 
scribed and figured separately with the hope that future collections of the /on- 
giflagrata complex will aid in their classification. 

Specimen no. |.—Preparatory female, body length 4.7 mm, RRH. ABYPL 
DS11, 42°59.7-59.2'N, 14°05.4-03.4’W, 5260 m. 

Description.—Frons and Rostrum (Fig. 6A—B): Anterior tip slightly overhang- 


466 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


ing frons. Rostrum length 0.13 cephalon width. Medial part of frons ridge slightly 
overhanging clypeal insertion. 

Antennula (Fig. 6D): Length 0.33 body length, with 13 flagellar articles. Distal 
5 annuli with single aesthetascs and 2 on tip. Article 2 length 0.68 medial length 
of article 1. Article 3 length 1.6 article 2 length. Medial lobe of article 1 lacking 
large setae or teeth. 

Left mandible (Fig. 6E): Spine row with 8 members. Molar process with 15 
setae on posterior margin of distal edge. Condyle length 0.16 mandibular body 
length. Palp article 2 length 0.54 mandibular body length. 

Maxilliped (Fig. 6F): Endite with 6 small and | large fan setae distally, and 4 
coupling hooks medially. Epipod length 0.97 basis length. 

Pleopod II (Fig. 6C): Length 0.28 depth. Apex distance to distal tip 0.36 total 
pleopod length. Keel somewhat rounded in lateral view. 

Uropod: Endopod length 1.5 protopod width; width 0.24 length. Exopod length 
0.82 endopod length. Protopod ventral surface with 7 large setae; dorsal surface 
with 2 setae near medial margin. 

Specimen no. 2.—Male, body length approximately 3.5 mm, RRH. INCAL 45, 
48°18.9-18.3'N, 15°14.4—13.3'W, 4829 m. 

Description.—Rostrum and frons similar to preparatory female (ABYPL DS11). 
Pleotelson lacking dorsal groove, with only slight indentation above uropod. 

Antennula (Fig. 7A): Article 2 length 0.8 medial length of article 1. Medial lobe 
of article | distal tip with 2 thick setae and 6 tiny denticles. 

Left mandible (Fig. 7B—D): Spine row with 6 members. Setal row on molar 
process with 11 members; triturating surface with 5 sensory pits. Condyle length 
0.18 mandibular body length. 

Maxilliped: Endite distal tip with 5 small and | large fan setae; medial margin 
with 3 coupling hooks. Epipod length 0.91 width. 

Pleopod I (Fig. 7E): Length 4.4 width at dorsal orifice. Distance from dorsal 
orifice to distal tip 0.30 total length. Ventral surface with pair of plumose setae 
proximally and 2 rows of fine setae distally. Lateral lobes similar to those in E. 
hessleri but longer; distally rounded with curved row of fine setae. Medial area 
of distal tip u-shaped with paired tufts of setae on inner edge. 

Pleopod II (Fig. 7F): Protopod length 1.9 width; lateral margin with 4 plumose 
setae. Endopod (including stylet) length 3.0 protopod length. 

Uropod (Fig. 7G): Endopod length 1.7 protopod width; width 0.23 length. Ex- 
opod length 0.76 width. Ventral surface of protopod with 4 large setae; medial 
edge of dorsal surface with | seta. 

Specimen no. 3.—Male fragment, cephalon missing, estimated body length 3.7 
mm, RRH. WHOI 330, see E. errabunda for station data. 

Description.—Pleopod I (Fig. 7H): Length 3.8 width; distance from distal tip 
to dorsal orifice 0.33 total length. Ventral surface and distal tip similar to male 
from INCAL 45 except: 2 plumose setae on left side and 4 plumose setae on 
right; medial area of distal tip with setae on small projections. 

Pleopod II: Protopod length 1.8 width; lateral margin with 2 plumose setae. 
Endopod length 3.1 protopod length. 

Remarks.—These individuals are most similar to Eurycope hessleri although 
the uropod of Eurycope sp. has much longer rami, and the antennular article 3 
is longer in the female. Differences in the pleopods from the two males (WHOI 


VOLUME 96, NUMBER 3 467 


330, INCAL 45) made identifying them as the same species uncertain. All three 
individuals occur in the same general area, the deep-sea adjacent to the Bay of 
Biscay, so they may possibly be the same species. The male from WHOI 330 was 
not classified as E. errabunda because it was estimated to be only 3.7 mm long 
and was fully adult; E. errabunda specimens are adult at much larger sizes, 
approximately 6 mm body length. 


Acknowledgments 


Sincere thanks are due the individuals and institutions mentioned in the Ma- 
terials and Methods section above for providing the specimens discussed in this 
paper. Helpful reviews of this paper by Robert R. Hessler, Scripps Institution of 
Oceanography and by Thomas E. Bowman and Brian Kensley, Smithsonian In- 
stitution, improved its content and form. This research was funded by grants 
from the National Science Foundation, DEB 77-24614 and DEB 80-07150, R. R. 
Hessler, principal investigator. 


Literature Cited 


Hansen, H. 1895. Isopoden, Cumaceen und Stomatopoden der Plankton-Expedition —Ergebnisse 
der Plankton-Expedition der Humboldt-Siftung 2:1—105. 

Kensley, B. 1977. New records of the marine Crustacea Isopoda from South Africa.—Annals of 
the South African Museum 72:239-265. 

Menzies, R. 1962. The isopods of abyssal depths in the Atlantic Ocean.—Abyssal Crustacea, Vema 

Research Series, Columbia University, New York 1:79-206. 

, and J. Pettit. 1956. A new genus and species of marine asellote isopod, Caecianiropsis 
psammophila, from California.—Proceedings of the United States National Museum 106:441- 
446. 

Nordenstam, A. 1933. Marine Isopoda of the families Seroloidae, Idotheidae, Pseudidotheidae, 
Arcturidae, Parasellidae and Stenetriidae mainly from the South Atlantic.—Further Zoological 
Results of the Swedish Antarctic Expedition 1901-1903, 3:1—284. 

Sars, G. 1864. Om en anomal Gruppe af Isopoder.—Forhandlinger 1 Videnskaps-Selskapet in Kris- 

tiana 1863:205—221. 

1899. Isopoda.—Crustacea of Norway 2:1—270, Bergen. 

Schultz, G. 1976. Species of asellotes (Isopoda: Paraselloidea) from. Anvers Island, Antarctica.— 
Antarctic Research Series 26: 1-35. 

Sivertsen, E., and L. Holthuis. 1980. The marine isopod crustacea of the Tristan da Cunha Archi- 
pelago.—Gunneria 35: 1-128. 

Wilson, G. 1976. The systematics and evolution of Haplomunna and its relatives (Isopoda, Hap- 
lomunnidae N. Fam.).—Journal of Natural History 10:569—580. 

——. 1981. Taxonomy and postmarsupial development of a dominant deep-sea eurycopid isopod 
(Crustacea).—Proceedings of the Biological Society of Washington 94:276—-294. 

———. 1983. Systematics of a species complex in the deep-sea genus Eurycope, with a revision of 

six previously described species (Crustacea, Isopoda, Eurycopidae).—Bulletin of the Scripps 

Institution of Oceanography 25: 1—64. 

, and R. Hessler. 1980. Taxonomic characters in the morphology of the genus Eurycope 

(Crustacea, Isopoda) with a redescription of E. cornuta Sars, 1864.—Cahiers de Biologie Ma- 

rine 21:241—263. 

, and 1981. A revision of the genus Eurycope (Isopoda, Asellota) with descriptions 

of three new genera.—Journal of Crustacean Biology 1:401—423. 

Wolff, T. 1962. The systematics and biology of bathyal and abyssal Isopoda Asellota.—Galathea 
Report 6: 1-320. 


A-002, Scripps Institution of Oceanography, La Jolla, California 92093, USA. 


PROC. BIOL. SOC. WASH. 
96(3), 1983, pp. 468-477 


SPONGE-INHABITING BARNACLES (CIRRIPEDIA: 
ARCHAEOBALANIDAE) OF THE CAROLINIAN 
PROVINCE, SOUTHEASTERN UNITED STATES, 
WITH THE DESCRIPTION OF A NEW SPECIES 

OF MEMBRANOBALANUS PILSBRY 


Victor A. Zullo and Jon D. Standing 


Abstract.—Acasta cyathus Darwin, 1854, previously known from the tropical 
western Atlantic north to Sapelo Island, Georgia, is reported from a variety of 
demosponges on the middle and outer continental shelf between Cape Fear and 
Cape Lookout, North Carolina. Membranobalanus declivis (Darwin, 1854), known 
previously from the tropical western Atlantic south of Cape Canaveral, Florida, 
occurs in Spheciospongia vesparium (Lamarck) from the mid-shelf region off 
Cape Fear, North Carolina. Membranobalanus costatus, new species, is found 
in Anthosigmella varians (Duchassaing and Michelotti) off Cape Fear. The new 
species is most readily distinguished from M. declivis by its prominently ribbed 
shell and rostrum of normal length. The number and distribution of specimens 
encountered in this study suggest that sponge-inhabiting barnacles are a common, 
but overlooked element of the Carolinian Province. 


Species of the archaeobalanid barnacle genera Acasta Leach and Membrano- 
balanus Pilsbry are obligate symbionts of sponges, occurring in tropical, sub- 
tropical, and warm temperate regions of the world. Most of the species ascribed 
to these genera are found in Indo-West Pacific waters, and only two species, one 
of each genus, have been reported from the western Atlantic. Acasta cyathus 
Darwin, 1854, is known from southern Georgia, southern Florida, the eastern 
Gulf of Mexico, and the Caribbean (Wells 1966; Newman and Ross 1976; Zullo 
and Lang 1978; Spivey 1981), but is also found in the eastern Atlantic, Red Sea, 
and Indo-west Pacific (Newman and Ross 1976). Membranobalanus declivis (Dar- 
win, 1854) has been reported from Bermuda, southern and western Florida, and 
the West Indies (Wells 1966; Newman and Ross 1976; Spivey 1981). 

The U.S. Bureau of Land Management (BLM) Live Bottom Study, conducted 
by the Duke University Marine Laboratory on the North Carolina continental 
shelf, has yielded an array of sponges, some of which contain sponge barnacles. 
Acasta cyathus is the most ubiquitous species, occurring in several different 
sponges at depths between 17 and 116 m. Membranobalanus declivis was dis- 
covered in two specimens of the loggerhead sponge, Spheciospongia vesparium 
(Lamarck), between 22 and 30 m depth southwest of Cape Lookout. Specimens 
of another sponge, Anthosigmella varians (Duchassaing and Michelotti), collect- 
ed off Cape Fear at depths between 28 and 32 m, contain a new and rather unusual 
species of Membranobalanus characterized by a prominently ribbed shell and a 
rostrum of normal length. The presence of Acasta and Membranobalanus on the 
North Carolina continental shelf extends the western Atlantic range of these 


VOLUME 96, NUMBER 3 469 


@Z-704 @O0S05 


CAPE FEAR 


@® MS04 


Fig. |. General location of stations for sponge-inhabiting barnacles off the North Carolina coast. 


genera nearly to the northern limit of the Carolinian Province. North Carolina 
localities are shown in Fig. 1. 


Station Descriptions 


1S05.—South-southwest of Cape Lookout, North Carolina, 34°23.0'N, 76°34.0’W, 
19-27 m, BLM Live Bottom Study station, 1981. 

MS04.—Southeast of Cape Fear, North Carolina, 33°31.0'N, 77°25.0’W, 27-55 
m, BLM Live Bottom Study station, 1981. 

OS05.—East of Cape Fear, North Carolina, 33°49.0’N, 76°33.5’W, 55-100 m, 
BLM Live Bottom Study station, 1981. 

Z-704.—WR-4 buoy, east-northeast of Cape Fear, North Carolina, 33°51.9'N, 
77°29.4'W, on superstructure of wreck, 17 m, Department of Earth Sciences 
collection, University of North Carolina at Wilmington, T. Prestia and P. 
Wheaton, collectors, 10 July 1982. 


470 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Systematic Account 


Subclass Cirripedia Burmeister, 1834 

Order Thoracica Darwin, 1854 
Suborder Balanomorpha Pilsbry, 1916 

Superfamily Balanoidea (Leach), Newman and Ross, 1976 
Family Archaeobalanidae Newman and Ross, 1976 
Subfamily Archaeobalaninae Newman and Ross, 1976 
Genus Acasta Leach, 1817 
Acasta cyathus Darwin, 1854 
Fig. 2a—e 


Acasta cyathus Darwin, 1854:312, pl. 9, figs. 3a—c (Madeira; West Indies).— 
Pilsbry, 1916:244, text-figs. 79-80, pl. 57, figs. 1-3 (Dry Tortugas, Florida; 
Colon, Panama; St. Croix, U.S. Virgin Islands).—Wells, 1966:85 (Gulf coast 
of Florida)—Newman and Ross, 1976:53 (Florida; Caribbean; Madeira; Mo- 
rocco; East Africa; Red Sea; Gulf of Manaar; Singapore; Kei Islands; Sulu 
Archipelago; Philippines; western Australia; 15-180 m; see for complete syn- 
onymy).—Zullo and Lang, 1978:159 (off Sapelo Island, Georgia).—Spivey, 1981: 
Wee 


Material examined.—Specimens of Acasta cyathus were obtained from the 
following localities and sponges off North Carolina: 


USNM hypotype lot Station Depth Date Host sponge 
USNM 195285 MS04 34 m 7 Feb 1981 Verongula ardis 
(de Laubenfels)? 
USNM 195286 OS05 66 m 3 Mar 1981 Family Coppatiidae sp. A 
USNM 195287 OS05 77 m 3 Mar 1981 Family Coppatiidae sp. A 
USNM 195288 OS05 62 m 3 Mar 1981 Cinachyra keukenthali 
Uliczka 
USNM 195289 OS05 69 m 4 Mar 1981 Family Coppatiidae sp. A 
USNM 195290 OS0S 64 m 4 Mar 1981 Erylus ministrongulus 
Hechtel 
USNM 195291 OS05 116 m 14 May 1981 Family Coppatiidae sp. A 
USNM 195292 OS05 104 m 14 May 1981 Family Coppatiidae sp. A 
USNM 195293 OS05 99 m 14 May 1981 Family Coppatiidae sp. A 
USNM 195294 MS04 32 m 18 May 1981 Family Mycalidae sp. A 
USNM 195295 MS04 30 m 10 May 1981 Ircinia campana (Lamarck) 
Z-704 17 m 10 Jul 1982 undetermined 


Remarks.—The 40 Acasta cyathus specimens from station Z-704 (from a single 
25 cm diameter, bowl-shaped sponge) represent a northern range extension of 
2°22’ in latitude and indicate that the species is common in North Carolina waters. 
The absence of prior records from this region is most likely the result of lack of 
collection, rather than a recent immigration from southern regions. 

The North Carolina Acasta were found in at least six different species of De- 
mospongea, occurring most frequently in an unidentified coppatiid sponge. Other 
host records for A. cyathus from the western Atlantic include sponges of the 
genus Aplysina (=Verongia), Ircinia campana (Lamarck), Ircinia felix (Duchas- 
saing and Michelotti) (=/. fasiculata), Spinosella (=Callyspongia) vaginalis (La- 
marck), and Spongia tubulifera Lamarck, (=S. officinalis) (Darwin 1854; Wells 
1966). 


VOLUME 96, NUMBER 3 47\ 


Comparison of North Carolina and Georgia specimens of Acasta cyathus re- 
vealed no important morphological differences between the two populations. 
Acusta cyathus is readily distinguished from other western Atlantic sponge bar- 
nacles by its pink-hued, globose shell, spinose parietes, small and nearly flat 
calcareous basis, obviously striate scutum lacking a depressor muscle pit, and 
narrow, rather long tergal spur. 


Membranobalanus Pilsbry, 1916 
Membranobalanus declivis (Darwin, 1854) 
Fig. 2f-o 


Balanus declivis Darwin, 1854:275, pl. 7, figs. 4a—d (West Indies; Jamaica).— 
Pilsbry, 1916:230, text-figs. 73-74, pl. 55, figs. 1-Id (off Cape Sable, Florida; 
Bermuda).—Pearse, 1932:119 (Dry Tortugas).—Wells, 1966:83 (Gulf coast of 
Florida). 

Balanus declivis var. cuspidatus Verrill, 1901:22 (Bermuda). 

Membranobalanus declivis (Darwin).—Newman and Ross, 1976:53 (see for syn- 
onymy).—Spivey, 1981:172. 


Material examined.—One individual in a specimen of Spheciospongia vespar- 
ium (Lamarck), off Cape Lookout, North Carolina, Hypotype USNM 195282, 
Station ISO5, 34°23.6'N, 76°34.9’W, 22 m, 9 Feb 1981; 5 individuals in a specimen 
of S. vesparium, off Cape Fear, North Carolina, Hypotype USNM 195283, Hy- 
potype lot USNM 195284, Station MS04, 33°31.4’N, 77°24.0’W, 30 m, 19 May 
1981. 

Remarks.—Yhe Membranobalanus declivis specimen from Station ISO5 ex- 
tends the northern range of this species by 9°15’ of latitude. 

Both Pearse (1932) and Wells (1966), who are the only authors to our knowledge 
that identified the host sponge for M. declivis, list the loggerhead sponge, Sphe- 
ciospongia vesparium. Their records, together with the new North Carolina rec- 
ords, suggest that S$. vesparium is the sole host for Membranobalanus declivis. 
The North Carolina M. declivis agree with the descriptions and illustrations pre- 
sented by Darwin (1854) and Pilsbry (1916). Membranobalanus declivis is distin- 
guished from Acasta cyathus by its membranous basis and non-spinose parietes, 
and from other Western Hemisphere species of Membranobalanus by its non- 
costate shell bearing narrow to broad radii and an elongate rostrum. 


Membranobalanus costatus, new species 
Figs. 2p—x, 3a—x 


Holotype.—Complete shell, opercular plates, and body, USNM. 

Dimensions of holotype.—Height, 8.8 mm; carinorostral diameter 10 mm; lat- 
eral diameter, 7.8 mm. 

Type locality. —Station MS04, 33°31.0'N, 77°25.0'W, off:Cape Fear, North Car- 
olina, 28 m depth, 12 Aug 1981. 

Material examined.—8 individuals from a specimen of Anthosigmella varians 
at the type-locality, Holotype USNM 195270, paratypes USNM 195271 through 
195277; 4 individuals from a specimen of A. varians, Station MS04, 33°32.8'N, 
77°24.3'W, 32 m, 18 May 1981, paratypes USNM 195278 through 195281. 

Diagnosis.—Shell thin, white, with radii lacking or incipient; orifice deeply 


472 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


N 
Sor 


te eee 


Fig. 2. Acasta cyathus Darwin, station Z-704: a, Shell, hypotype USNM 195285; b—e, Outer and 
inner views of opercular plates, hypotype USNM 195285. Membranobalanus declivis (Darwin), sta- 
tion MS04: f-m, Outer and inner views of opercular plates, hypotype USNM 195282; n—o, Side and 
exterior view of rostrum, hypotype USNM 195282. Mouth parts and cirri of M. costatus n. sp., 
paratype USNM 195278, station MS04: p, Palp; q, Second maxillae; r, First maxilla; s, Labrum; t, 
Mandible; u, Cirrus I; v, Cirrus II; w, Cirrus VI; x, Medial articles of Cirrus VI. Scale: Fig. a, <3; 
figs. b-m, <6; figs. n-o, x4; figs. p—q, x20; figs. u-v, x10; figs. w—x, x20. 


VOLUME 96, NUMBER 3 473 


toothed; rostrum same length as other compartmental plates; exterior of parietes 
with narrow, prominent, irregularly-spaced costae; opercular plates without in- 
tercalated chitinous lamellae; scutal adductor ridge incipient or lacking; tergal 
spur short, broad, usually occupying two-thirds of basal margin; outer ramus of 
Cirrus IV with up to 6 recurved teeth per article; medial articles of Cirrus VI 
with 3 pairs of spines per article, proximal pair minute. 

Description.—Shell (Fig. 3a—j) high-conic, white, with deeply-toothed orifice; 
compartmental plates solid, thin, easily disarticulated; parietes ornamented ex- 
ternally with fine, closely-spaced, irregular growth lines crossed by fine, longi- 
tudinal striae on upper (younger) third and by prominent, narrow, irregularly- 
spaced costae on lower two-thirds; costae extend slightly below basal margin of 
parietes giving base of shell wall a denticulate appearance; rostrum larger than 
other compartmental plates, but not of greater length, narrow in upper third and 
increasing in width towards broadly V-shaped basal margin; carina narrow with 
nearly parallel parietal borders; apices of rostra and carinae with deeply incised 
notches, probably caused by cirral rasping; radii absent or at best represented by 
extremely narrow and irregular ledges; alae narrow and steeply oblique in upper 
third of compartmental plate, broadening markedly in central third, and narrowing 
abruptly in lower third; alar sutural edges smooth; sheath of rostrum one-half 
length of plate; sheath of other compartmental plates one-half to two-thirds length 
of plate; lower margin of sheath continuous with shell wall; interior of parietes 
often ribbed below sheath; internal ribs obverse of sulcations between external 
costae; basis membranous. 

Scutum (Fig. 3k—n, u—x) thick, markedly convex, white, with adherent exterior 
yellow-brown epidermis bearing fine hairs; basal margin shorter than tergal mar- 
gin; tergal margin reflexed about 45°; exterior ornamented by narrow, closely- 
spaced, semi-erect growth ridges crossed by very fine radial striae, especially 
evident in medial part of plate, every other external growth ridge forming raised 
‘“‘tooth’’ on occludent margin; articular ridge prominent, triangular, between three- 
fifths and three-fourths length of tergal margin, and reflexed over narrow, shallow 
articular furrow; reflexed articular ridge extends well beyond tergal margin of 
scutum; adductor ridge represented by low, rounded, raised border of small, 
deep, oval adductor muscle pit just above and tergad of center of plate; depressor 
muscle pit large, deep, triangular, extending up under inner lamina of scutum and 
located at basal margin in angle formed by reflexed tergal margin; narrow, tri- 
angular ‘“‘radius,’’ broadest at apical end, extending length of occludent margin. 

Tergum (Fig. 30—t) thin, about same width as scutum, white; basal margin 
slightly shorter or equal to length of scutal margin; exterior ornamented by fine, 
closely-spaced growth ridges, and covered in lower half by yellow-brown epi- 
dermis bearing fine hairs; spur furrow broad, very slightly depressed, bounded 
on either side by impressed lines; narrow strip along carinal border of tergum 
depressed below exterior surface of plate and with upturned growth lines; tergal 
spur short, broad (base of spur occupying about two-thirds of basal margin), 
basally truncate at angle to basal margin, and placed close to basiscutal angle; 
length of spur about one-half basal width of spur; depressor muscle crests absent, 
or represented by few inconspicuous, irregular ridges; articular ridge straight, 
erect, short, restricted to apical half of plate; articular furrow very broad, shallow. 


474 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 3. Membranobalanus costatus n. sp., station MS04: a, Shell, holotype USNM 195270; b, 
Shell, paratype USNM 195271; c—d, Outer and inner view of rostrum, paratype USNM 195273; e-f, 
Outer and inner views of lateral, same specimen; g—h, Outer and inner views of carinolateral, same 
specimen; i-j, Outer and inner views of carina, same specimen; k—n, Outer and inner views of scuta, 
paratype USNM 194274; o—r, Outer and inner views of terga, same specimen; s—t, Outer and inner 
views of tergum, paratype USNM 195275; u—x, Outer and inner views of scuta, same specimen. 
Scale: Figs. a—b, x4; figs. c—j, <7; figs. k-x, x8. 


VOLUME 96, NUMBER 3 475 


Labrum (Fig. 2s) triangular, with deep notch at apex of crest; two to six teeth 
present on one or both sides of notch. 

Palp (Fig. 2p) kidney-shaped, superior margin concave, densely setose, setae 
short, pectinate; inner margin densely setose, setae longer, pectinate; inferior 
margin nearly devoid of setae. 

Mandible (Fig. 2t) with fine teeth on cutting edge; upper tooth largest; second 
tooth not bifid, slightly smaller than first tooth, and located at center of cutting 
edge; third tooth slightly smaller than second, located at center of lower half of 
cutting edge; fourth and fifth teeth considerably smaller, fourth larger than fifth, 
fifth barely visible in some specimens; inferior angle without obvious denticles 
Or spines (may be result of wear); inferior and superior margins setose. 

Maxilla I (Fig. 2r) with straight edge; uppermost 2 large spines of equal size; 
notch lacking or incipient below uppermost 2 spines, its position indicated by | 
or 2 minute spinules; middle section bears 4 or 5 shorter, more slender, and non- 
alternating spines; lower third of edge bears 4 or 5 spines of same size as upper- 
most pair; inferior angle with tuft of small spinules; inferior and superior margins 
setose, with those of inferior margin longer, more numerous. 

Maxilla II (Fig. 2q) bilobed; outer lobe ovate, superior and posterior margins 
densely setose, setae of posterior margin primarily non-pectinate, those of su- 
perior margin pectinate; inner lobe small, circular, densely clothed with coarsely 
pectinate setae. 

Cirrus I (Fig. 2u) with grossly unequal rami; anterior ramus about 3 times length 
of posterior ramus, anterior ramus reversed, antenniform; posterior ramus more 
setose than anterior, with slightly protuberant articles; Cirrus II (Fig. 2v) short, 
with outer ramus one-third again as long as inner ramus; articles of both rami 
slightly protuberant, covered with pectinate setae somewhat longer than those of 
Cirrus I, outer distal margins of articles bear comb-like patches of minute spinules 
at bases of setae; Cirrus III longer and more slender that Cirrus II; outer ramus 
longer than inner; articles of rami not protuberant; setation and distribution of 
spinule comb rows similar to those of Cirrus II; Cirri IV—VI longer and more 
slender than Cirri I-III, gradually increasing in length posteriorly; Cirrus IV (Fig. 
2v) with inner ramus slightly longer than outer; pedicel about one-half length of 
outer ramus; outer ramus less slender than inner, bearing large, recurved teeth, 
up to 6 in number, on medial articles; distal margins of outer surface of pedicel 
and articles each bearing row of upright denticles; inner ramus with few upright 
denticles at distal-anterior corner, and comb-like rows of minute spinules along 
outer distal borders of articles; Cirrus V longer and more slender than Cirrus IV, 
with denticles and comb-like rows of spinules on both rami arranged as on inner 
ramus of Cirrus IV Cirrus VI (Fig. 2x) longer than Cirrus V, with 3 pairs of spines 
on medial articles, distal pair long, median pair short, proximal hair minute; distal 
margins of outer surface of articles with few comb-like patches of spinules. 

Etymology.—The specific name costatus is Latin for ribbed or costate, and 
refers to the externally costate shell of the new species. 

Discussion.—Membranobalanus costatus is readily distinguished from previ- 
ously described species of Membranobalanus by its costate parietes. The new 
species is related to Western Hemisphere Membranobalanus, all of which bear 
large, recurved teeth on the outer ramus of Cirrus IV. Of these species, M. 


476 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


costatus 1S most similar in appearance to M. nebrias (Zullo and Beach, 1973) 
from the Galapagos Archipelago. In both species the shells lack radii, the rostra 
are short, and the opercular plates, trophi, and cirri are similar in overall con- 
formation. Membranobalanus costatus differs consistently from M. nebrias in 
several features: (1) the rostrum is of ‘“‘normal’’ length; (2) the opercular plates 
lack intercalated chitinous lamellae; (3) the scutum lacks a rostral depressor mus- 
cle pit in the basioccludent angle; (4) the scutal articular ridge is longer; (5) a 
‘“‘radius’’ 1s developed along the occludent margin of the scutum; (6) the tergal 
spur is broader; (7) the tergal articular ridge is longer; (8) the outer lobe of Maxilla 
II is much shorter; (9) the outer ramus of Cirrus IV bears up to six, rather than 
four, recurved teeth per article; and (10) the medial articles of Cirrus VI have up 
to three, rather than five, pairs of spines. 

Membranobalanus costatus also resembles M. declivis, especially in features 
of the trophi and cirri and in the possession of a “‘radius’’ along the occludent 
margin of the scutum. Membranobalanus costatus is distinguished by its much 
shorter rostrum, absence of radii, shorter and thicker scutum that is markedly 
bowed from side to side, rather than nearly flat, its shorter and more equally 
triangular tergum, shorter and broader tergal spur, and vertical rather than oblique 
articular ridge. 

Membranobalanus costatus differs from M. orcutti in having a short rostrum, 
a broader scutum with a much shorter adductor ridge and a “‘‘radius’’ along the 
occludent margin, a narrower tergum with a longer tergal spur, a simpler armature 
on the outer ramus of Cirrus IV, and three, rather than four, pairs of spines on 
the medial articles of Cirrus VI. The new species is most readily distinguished 
from the Indo-West Pacific species M. brachialis (Rosell), M. cuneiformis (Hiro), 
and M. longirostrum (Hoek) by the presence of large, recurved teeth on Cir- 
rus IV. 

If M. costatus is specific to Anthosigmella varians, as M. declivis appears to 
be with its host sponge, then this new barnacle species might be expected to 
occur in Florida, the Gulf of Mexico, the West Indies, and the Caribbean side of 
Central America (cf. Wiedenmayer 1977:245). We hope that queries such as this 
will stimulate further research on this interesting group of symbiotic barnacles. 


Key to Western Hemisphere Species of Acasta and Membranobalanus 


1. Basis membranous; shell high-conic, without calcerous spines on exterior 


OL PANELS Si eae sc sh a ond. cal pene ons dee 2 oe ecient ar 2 
— Basis calcareous, slightly cup-shaped; shell globular, with calcareous spines 
ONLEXfeCMOr Ob PaNletes: ny. cugees «oe y 6s ee ome Acasta cyathus 
2. Rostrum considerably longer than other compartmental plates, usually at 
least.twice length of carina... . 6. ci-cgoee 0S0ee ce eo ee eee 3 
— Rostrum slightly longer or not noticeably longer than other compartmental 
PLALSS ace 2's sheaf eee AN ee ol athe Sete SA 4 


3. Radii present, narrow to broad; basal margin of scutum about same length 
as tergal margin; scutal adductor ridge, if present, confined to adductor 
muscle pit border; occludent margin of scutum bordered by sunken ledge 
CLT ACU) Re ae saa wets ata, eee aie ee Membranobalanus declivis 
— Radi lacking; basal margin of scutum considerable shorter than tergal 


VOLUME 96, NUMBER 3 477 


margin; scutal adductor ridge prominent, extending to basal margin; oc- 
cludent margin of scutum lacks accessory ledge ... Membranobalanus orcutti 
4. Exterior of parietes smooth, except for growth bands; chitinous lamellae 
intercalated with calcareous lamellae of opercular plate apices; occludent 
margin of scutum lacks accessory ledge ........ Membranobalanus nebrias 
— Exterior of parietes prominently costate; apices of opercular plates lack 
chitinous lamellae; occludent margin of scutum bordered by sunken ledge 
CO°PAGIOS OY PoP Ra ene eee Membranobalanus costatus 


Acknowledgments 


We thank Kirstin Sandoy for first calling our attention to the sponge barnacles 
in the BLM material; William Kirby-Smith for allowing us access to the speci- 
mens; F. L. Nicholson and Kirstin Sandoy for identifying the sponges; and Thom- 
as Prestia and Patricia Wheaton for donation of Acasta cyathus specimens from 
locality Z-704. This study was funded by the Marine Sciences Program, Univer- 
sity of North Carolina at Wilmington, the National Science Foundation (Grant 
no. DEB-8020379), and the Crustacean Research Fund. 


Literature Cited 


Darwin, C. 1854. A monograph on the subclass Cirripedia with figures of all the species. The 
Balanidae, the Verrucidae, etc.—Ray Society, London, 684 pp. 

Newman, W. A., and A. Ross. 1976. Revision of the balanomorph barnacles; including a catalog 
of the species.—San Diego Society of Natural History, Memoir 9, 108 pp. 

Pearse, A. S. 1932. VII. Inhabitants of certain sponges at Dry Tortugas. Papers from Tortugas 
Laboratory.—Carnegie Institution of Washington (Publ. 435) 28:117-124. 

Pilsbry, H. A. 1916. The sessile barnacles (Cirripedia) contained in the collections of the U.S. 
National Museum; including a monograph of the American species.—Bulletin of the United 
National Museum 93: 1-366. 

Spivey, H.R. 1981. Origins, distribution, and zoogeographic affinities of the Cirripedia (Crustacea) 
of the Gulf of Mexico.—Journal of Biogeography 8:153-176. 

Vernll, A. E. 1901. II. Additions to the fauna of the Bermudas from the Yale Expedition of 1901, 
with notes on other species.—Transactions of the Connecticut Academy of Arts and Sciences 
11:15-62. 

Wells, H. W. 1966. Barnacles of the northeastern Gulf of Mexico.—Quarterly Journal of the Florida 
Academy of Sciences 29:81-95S. 

Wiedenmayer, F. 1977. Shallow-water sponges of the western Bahamas.—Birkhauser, Basel and 
Stuttgart, 287 pp. 

Zullo, V. A., and D. B. Beach. 1973. New species of Membranobalanus Hoek and Hexacreusia 
Zullo (Cirripedia, Balanidae) from the Galapagos Archipelago.—Los Angeles County Natural 
History Museum Contributions in Science 249: 1-16. 

Zullo, V. A., and W. H. Lang. 1978. Subclass Cirripedia. Pp. 158-160 in R. G. Zingmark, ed., An 
annotated checklist of the biota of the coastal zone of South Carolina.—University of South 
Carolina Press, Columbia, 364 pp. 


(VAZ) Department of Earth Sciences, University of North Carolina at Wil- 
mington, Wilmington, North Carolina 28403; (JDS) Duke University Marine Lab- 
oratory, Beaufort, North Carolina 28516; present address: Bodega Marine Lab- 
oratory, Bodega Bay, California 94923. 


PROC. BIOL. SOC. WASH. 
96(3), 1983, pp. 478-480 


ZEUGOPHILOMEDES, A NEW GENUS OF MYODOCOPINE 
OSTRACODE (PHILOMEDINAE) 


Louis S. Kornicker 


Abstract.—Zeugophilomedes, a new genus of myodocopine ostracode in the 
subfamily Philomedinae is proposed for 5 species in the Red Sea and Indian, 
Atlantic and Pacific Oceans. A key to the species is given and pertinent morpho- 
logical characters illustrated. 


A new genus Is proposed for 5 species previously referred to either Philomedes 
Liljeborg, 1853 or Euphilomedes Poulsen, 1962: Zeugophilomedes oblongus (Ju- 
day, 1907:145), Z. polae (Graf, 1931:5), Z. multichelatus (Kornicker, 1958:230), 
Z. fonsecensis (Hartmann, 1959:197), and Z. arostratus (Kornicker, 1967a:2). 
Euphilomedes graft (Hartmann, 1964:37) may, when more completely known, 
be referred to Zeugophilomedes. 


Zeugophilomedes, new genus 
Figs. 1-3 


Type-species.—Philomedes multichelata Kornicker, 1958. 

Etymology.—The name derived by combining the Greek zewgos (=pair, team) 
and Philomedes. Gender: masculine. 

Diagnosis.—Sixth limb: End joint with relatively slight posterior projection. 

Furca (Figs. 1-3): Each limb with 4 or 5 primary claws: 2 anterior claws sep- 
arated from lamella by suture; remaining claws fused to lamella. One or more 
secondary claws between primary claws 2 and 3, and 3 and 4. Primary claw 4 
followed by minute primary claw or additional secondary claws. 

Internal sclerites in posterior part of body (Figs. 1-3): Y-sclerite unbranching, 
fairly stout. Complex of sclerites connected to proximal end of Y-sclerite. (Scler- 
ites are visible through body in transmitted light.) 

Comparisons.—Other genera in the Philomedinae have 6th limbs with end joints 
having considerable posterior projection. The slight posterior projection of the 
end joint of 6th limbs of Zeugophilomedes resembles that of members of the 
Pseudophilomedinae. I consider this to be the result of parallelism or conver- 
gence. (Because the 6th limb of Z. oblongus and Z. fonsecensis had not been 
described, I examined a female of the former [USNM 139159] from Monterey 
Bay, California, and an A-1 female of the latter from El Salvador and found the 
6th limbs to have only slight posterior projection.) The furcae of members of 
Zeugophilomedes are unique for the Philomedidae in having some primary claws 
fused to the lamella (Figs. I-3). The Y-sclerites (see Kornicker 1975:684 for 
discussion) of species in other genera of Philomedinae branch proximally (Y- 
Shaped), and the combined stem and dorsal branch is concave dorsally; it is not 
linear and unbranched as in Zeugophilomedes (Figs. 1-3). The complex of scler- 
ites of the type connected to the proximal end of the Y-sclerite of Zeugophi- 
lomedes (Figs. 1-3) has not been described in other genera. 


VOLUME 96, NUMBER 3 479 


Figs. 1-3. Posterior of body of 3 species of Zeugophilomedes showing furca and internal sclerites 
(arrow indicates Y-sclerite): 1,Z. arostratus 6, USNM 112659; 2, Z. multichelatus 6, USNM 152449; 
3, Z. oblongus 2, USNM 139159. 


Known distribution.—Z. oblongus, San Diego Bay and off San Pedro, Califor- 
nia (Juday 1907:147), Monterey Bay, California (herein). Z. polae, Gulf of Suez, 
Red Sea (Graf 1931:38; Kornicker 1967c:4). Z. multichelatus, Bimini, Bahamas 
(Kornicker 1958:232, 1967b:2), Aransas Pass, Gulf of Mexico (herein). Z. aro- 
stratus, Maldive Islands, Indian Ocean (Kornicker 1967a:14). Z. fonsecensis, El 
Salvador (Hartmann 1959:198). 


480 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Key to the Species of Zeugophilomedes 


1. Furca with minute primary claw following the 4th primary claw ....... D 
— Furca with | or more secondary claws following 4th primary claw ..... 3 
2. Furca with | secondary claw following 2nd primary claw ............. 


RE ER errr tyes Seen Cone er TR, be A Te 3 Ba Re Bl Z. fonsecensis 
— Furca with 2 or 3 secondary claws following 2nd primary claw ........ 


— Furca with 4 or 5 secondary claws following 2nd primary claw ............ 
icp abe Neen npr pass SCR Ee Me tae. 6 Oa MERON Z. multichelatus 
3. Incisur of female carapace very shallow; 2nd endopodial joint of male 2nd 


antenna with 2 subtermimeali bristles! 4.55 4-5 504 ee ome ee eee Z. polae 
— Incisur of female carapace fairly deep; 2nd endopodial joint of male 2nd 
antennavwith 2 mudbristles: s.< a6 «2-year Z. arostratus 
Acknowledgments 


I thank the following people for their help: Gerd Hartmann for loan of a spec- 
imen of Z. fonsecensis from El Salvador; Peter M. Slattery for gift of a specimen 
of Z. oblongus from Monterey Bay, California; Carolyn Gast for inking illustra- 
tions; and Anne C. Cohen and Thomas E. Bowman for comments on the manu- 
script. 


Literature Cited 


Graf, H. 1931. Expedition S.M.S. “‘Pola”’ in das Rote Meer: Die Cypridinae des Roten Meeres.— 
Denkschriften der Akademie der Wissenschaften in Wien, Mathematsch, Naturwissenschaft- 
liche Klasse 102:32—46. 

Hartmann, Gerd. 1959. Zur Kenntnis der lotischen Lebensbereiche der pazifischen Kuste von El 
Salvador unter besonderer Berticksichtigung seiner Ostracodenfauna.—Kieler Meeresfor- 
schungen 15(2):187—241. 

—. 1964. Zur Kenntnis der Ostracoden des Roten Meeres.—Kieler Meeresforschungen, Son- 
derheft 20:35-127. 

Juday, Chauncy. 1907. Ostracoda of the San Diego Region, II. Littoral forms.—University of 
California Publications in Zoology 3(9): 135-156. 

Kornicker, Louis S. 1958. Ecology and taxonomy of Recent marine ostracodes in the Bimini area, 
Great Bahama Bank.—Publications of the Institute of Marine Science (The University of Tex- 
as) 5:194—300. 

——. 1967a. Euphilomedes arostrata, A new myodocopid ostracod from Maldive Islands, Indian 
Ocean.—Proceedings of the United States National Museum 120(3563): 1-21. 

———. 1967b. Supplementary description of the myodocopid ostracod Euphilomedes multichelata 
from the Great Bahama Bank.—Proceedings of the United States National Museum 120(3566): 
1-16. 

——. 1967c. Supplementary descriptions of two myodocopid ostracods from the Red Sea.— 
Proceedings of the United States National Museum 121(3571):1—-18. 

——. 1975. Antarctic Ostracoda (Myodocopina).—Smithsonian Contributions to Zoology 163:1— 
720 p. 

Liljeborg, Wilh. 1853. Ostracoda. Pp. 92-130 in De Crustaceis ex ordinibus tribus: Cladocera, 
Ostracoda et Copepoda in Scania Occurrenibus. 

Poulsen, E. M. 1962. Ostracoda-Myodocopa, 1: Cypridiformes-Cypridinidae.—Dana-Report 57:1— 
414. Copenhagen: Carlsberg Foundation. 


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


PROC. BIOL. SOC. WASH. 
96(3), 1983, pp. 481-488 


A NEW SPECIES OF MUNIDOPSIS FROM SUBMARINE 
THERMAL VENTS OF THE EAST PACIFIC RISE 
AT 21°N (ANOMURA: GALATHEIDAE)! 


Austin B. Williams and Cindy Lee Van Dover 


Abstract.—A new galatheid, Munidopsis lentigo, is described from depths of 
2600 m on a thermally active area of the East Pacific Rise. The species has 
distinctive organs on the chelae and strikingly flattened eyes. Similarities to other 
species in the genus are discussed. 


Exploration of thermally heated vent areas on rift zones of the East Pacific 
Rise has yielded yet another previously unknown decapod crustacean species 
which is here described. One of us (CLVD) participated in the OASIS Expedition 
to vent sites at 21°N aboard the R/V Melville, 12 April to 8 May 1982, during 
which collections of Munidopsis and other decapods were made by various in- 
vestigators with the aid of the deep submersible ALVIN on Dives 1211 and 1221. 


Munidopsis lentigo, new species 
Figs. 1-3 


Diagnosis.—Differs from all other species of Munidopsis in possession of len- 
ticular-shaped, flattened, smooth, light brown spot on ventral surface of each 
chela, and flattened eyes with depressed cornea shielded by projecting flat dorsal 
spine and ventral plate. 

Description.—Carapace, exclusive of rostrum, distinctly longer than broad, 
moderately arched transversely; cervical and transverse grooves faintly indicated, 
slight depression in cardiac region, scattered obsolescent rugosities on each an- 
terior branchial region, more distinct and transversely arranged rugosities on each 
posterior branchial region; posterior margin deeply, sometimes angularly emar- 
ginate medially. Frontal margin armed with strong triangular spine lateral to eye; 
anterior 74 of lateral margin armed typically with 7 spines. Rostrum slender, 
dorsally flattened, slightly deflexed and with serrate borders anteriorly, tip acute 
(bent to left in holotype). Lateral plate lightly rugose, projecting anteriorly below 
antennal peduncle and tipped anteriorly with obscure serration. 

Abdomen unarmed, second and third segments bearing low dorsal transverse 
ridge paralleled by shorter ridge bordering posterior margin; obsolescent anterior 
transverse ridge on fourth segment; fifth and sixth segments smooth. 

Eyes large, dorsoventrally flattened. Cornea cupped within overgrowths of 
ocular peduncle consisting of flat, broadly triangular, projecting dorsal spine with 
obscurely serrate margins having distal % slightly upturned and bent mesad to 
reach about % length of rostrum, continuous at each side with spatulate ventral 
lip serrated on margin. 


1 OASIS Expedition Contribution No. 5. 


482 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Basal article of antennular peduncle with single outer terminal spine; preceded 
by slender spine on dorsal margin, a very short terminal spine on mesial margin. 
Antennal peduncle with 2 fixed spines on basal article, lateral one spatulate and 
acutely tipped, ventral one slender; short second and fourth articles each bearing 
short buttressed lateral spine, much larger third article with serrated lateral flange. 

Third maxilliped with ischium 4% longer than merus and bearing mesial ridge 
armed with finely uniform, evenly spaced teeth; merus with distolaterally directed 
strong spine at midlength of ventrolateral border; carpus densely setose mesially, 
propodus and dactyl so on ventromesial borders. Projecting third thoracic somite 
with anterior margin nearly straight, lacking spines or median notch. 

Epipods absent from pereopods. 

Chelipeds of mature male asymmetrical, rugose proximally but becoming 
smoother distally, spined and sparsely but inconspicuously setose; cutting edges 
of fingers placed toward dorsal side. Major (right) cheliped of holotype robust; 
ischium unarmed; merus with very strong subdistal spine preceded by smaller 
spine on mesial margin, distolateral spine small and acute, rounded mesioventral 
ridge bearing acute spine near midlength and terminating in blunt distal spine; 
carpus with 2 strong spines on mesial margin; chela broad, obscurely rugose, 
palm as long as fingers, convex irregular lateral margin bearing 4 remote dorsal 
spines, similar mesial margin bearing 5 obsolescent spines, swollen ventral surface 
marked with 3 longitudinal lines of remote setal tufts on obsolescent ridges, mid- 
dle ridge becoming pronounced rib on fixed finger and curving mesad to terminate 
in hooked tooth; shallow concavity near base of fixed finger bearing slightly 
raised, flattened, smooth, light brown, bean-shaped spot (1.28 x 2.11 mm); fin- 
gers stout, more or less flattened above; cutting edges crenate, closing closely; 
fixed finger with single basal tooth fitting into notch distal to single basal tooth 
on dactyl, terminating in small dorsal tooth and slightly hooked ventral tooth; 
dactyl ending in hooked tooth confluent ventrally with truncate accessory cusp 
on rib running length of finger. Minor (left) cheliped of holotype acutely spined 
and sparsely but conspicuously setose; ischium unarmed; merus somewhat pris- 
matic, dorsal crest bearing row of small spines, 2 strong spines on mesial margin 
distally plus a smaller distoventral and still smaller distolateral spine; carpus with 
4 mesial spines grading from small proximally to very strong distally; chela some- 
what broadened, more or less flattened above but palm with fixed finger and dactyl 
independently somewhat trigonal below; slight convexity on lateral margin at base 
of fixed finger adjacent to ventral, slightly raised, flattened, smooth, light brown 
oval spot (1.34 x 1.98 mm); palm slightly longer than fingers, 5 or 6 spines on 
lateral margin, 4 less outstanding spines on mesial margin; fingers nearly straight, 
cutting edges closing closely, straight and indistinctly crenate, hooked tip of dac- 
tyl closing between hooked acute upper and broader based lower terminal teeth 
of fixed finger. 

Female with chelipeds analogous to minor cheliped of male; those of paratype 
191161 slightly asymmetrical and rather slender, ventral smooth spot on palm at 
base of fixed finger conspicuous (on right chela 0.54 x 0.99 mm, left 0.64 x 1.22 
mm); those of paratype 191162 also slightly asymmetrical but with chelae broader 
(fingers broken left side), ventral spot on palm at base of fixed finger (on right 
chela 1.02 x 1.34, left 0.96 x 1.34 [est.] mm). 


VOLUME 96, NUMBER 3 483 


First walking legs reaching to or beyond base of dactyl on cheliped, second 
and third legs reaching about to base of dactyl on preceding legs. Merus of each 
walking leg slender; row of small spines dorsally, strongest and most numerous 
on third, less developed on second and first; first and second with strong disto- 
lateral spine. Carpi armed with distodorsal spine and smaller distoventral spine. 
Propodi compressed, slightly shorter than meri, each bearing 2 or 3 slender mov- 
able spines ventrally and unequal pair of slender movable spines distoventrally 
near articulation of dactyl. Dactyls about 4 length of propodi, compressed, curved, 
corneous tip preceded by comb of anteriorly directed spinules on ventral margin. 


Measurements in mm.— 


M M F F F 
Holotype Paratype Paratype Paratype Paratype 
191160 191163 191163 191162 191161 
Carapace 
Base ocular peduncles 
to notch in 
posterior margin 12.8 10.6 8.2 11.5 11.9 
Rostrum 3.8 3.1 2.6 3o2) 3.5 
Total length 16.6 13.7 10.8 14.7 15.4 
Width 10.9 8.7 8.8 9.5 10.4 
Chelae 
Major 
Length propodus 16.6(R) missing 12.8(R) 12.1(L) 
Width propodus 6.8 5.0 3.1 
Length dactyl Ved 325) 6.1 
Minor 
Length propodus 16.6(L) missing *11.1(L) 10.5(R) 
Width propodus 4.4 4.1 Doll 
Length dactyl Toll *4.5 5.3 
Eggs 2.00 x 2.24 x 
2.20 2.24 


* Broken (estimated). 


Variation.—The posterior border of the carapace varies from evenly concave 
(usual) to notched medially (paratype F 191161). The latter specimen also has a 
more prominent and distinctly outlined cardiac region than do other members of 
the type-series, and the ventral margin of lateral plates on its cephalothorax is 
more rounded. Variability in chelae is noted in the description. 

Type-locality.—Pacific Ocean, East Pacific Rise, 20°49.6’N, 109°6’W, 2600 m. 

Material studied.—Confined to the type-series listed under measurements and 
deposited in the crustacean collection of the United States National Museum of 
Natural History, Smithsonian Institution (USNM), Washington, D.C. 

Etymology.—The name is a noun in apposition from the Latin “‘lentigo,’’ mean- 
ing a lentil-shaped spot, referring to the oval spot on the ventral side of each 
chela. 

Remarks.—Munidopsis lentigo has features that set it well apart from other 
species in the richly diverse genus Munidopsis, but a more comprehensive study 
than we have made should precede any attempts to separate it from that genus 


484 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. |. Munidopsis lentigo. Male holotype USNM 191160: a, Dorsal view (tip of rostrum bent to 
left), appendages not shown; b, Right chela, carpus, and distal part of merus. Scale = 2 mm. 


(see Chace 1942; Ambler 1980). No other known members of Munidopsis possess 
the peculiar oval organs on the chelipeds nor do any have flattened eyes with 
depressed cornea covered by a projecting flat dorsal spine as well as shielded by 
a ventral plate. Several species, however, have eyes armed with spines. Among 
those that can be compared with M. lentigo in this respect, M. beringana Ben- 
edict, 1902, M. ciliata’: Wood-Mason, 1891, M. crassa Smith, 1885, M. pilosa 
Henderson, 1885 (illustrated 1888), and M. verrilli Benedict, 1902 each have a 
slender compressed, somewhat upturned rostrum which leaves the eyestalks ex- 
posed to dorsal view. In all of these species except M. pilosa, the eyestalk is 
drawn into a prominent mesial spine exceeding the subglobular lateral cornea, 
and in M. ciliata and M. verrilli there is a small posterolateral spine as well. The 
eyestalk of M. pilosa terminates in a long slender spine that covers only the 
middorsal part of the subglobular cornea and reaches about half the length of the 
rostrum beyond it; there is also a short acute ventral spine but no ventral plate. 


VOLUME 96, NUMBER 3 485 


Fig. 2. Munidopsis lentigo. Female paratype USNM 191161: a, Dorsal view; b, Rostrum and eyes; 
c, Carapace, eye and base of antenna, lateral view; Basal article of antennal peduncle, distal part; d, 
Ventral view; e, Same, lateral view; f, Left cheliped, ventral view. Scales = 1 mm. 


486 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 3. Munidopsis lentigo. Female paratype 191162: a, Dorsal view, appendages not shown; b, 
Rostrum and eyes; c, Carapace, eye and base of antenna, lateral view; d, Rostrum and eyes, lateral 
view; e, Basal article of antennal peduncle, distal part, dorsal view; f, Same, lateral view; g, Right 


cheliped, dorsal view; h, Same ventral view; i, Carpus, left cheliped; j, Right second pereopod; k, 
Right third pereopod; |, Left fifth pereopod. Scales = | mm. 


VOLUME 96, NUMBER 3 487 


Other species such as M. bermudezi Chace, 1942, have eyestalks resembling those 
of species mentioned above, but the rostrum is broader at its base, while species 
such as M. nitida (A. Milne Edwards, 1880, illustrated by Milne Edwards and 
Bouvier, 1897) and M. scabra Faxon, 1895, have eyestalks with short, extremely 
slender mesiodorsal spines which hardly conceal the prominent globular cornea 
at all. The broad rostrum of M. /atirostris Faxon, 1895, is emarginate basally, 
exposing fixed eyestalks with subglobular cornea partly covered by a broad, 
mesiodorsal outgrowth of the peduncle. 

The oval spots on the chelipeds recall similar organs in homolid crabs of the 
genus Hypsophrys both in shape and placement at the base of the fixed finger 
(Williams 1974, 1976). Species of Hypsophrys displays a spot on both inner and 
outer surface of the chelae whereas M. lentigo has them only on the ventral 
surface, analogous to the inner surfaces in Hypsophrys. Fingers on the right chela 
of female 191162 were broken by an accidentally dropped camera lens during 
study, the break on the fixed finger passing through the spot on that hand. In- 
spection of the broken spot under a binocular dissecting microscope indicates an 
internal structure resembling that in the organs on A. nour Williams, 1974, and 
H. superciliosa Wood-Mason, 1891, whose function is unknown (Williams 1976). 
Histological structure of the spots in M. /entigo remains to be studied. 

In none of the comparable species of Munidopsis does armature of the integ- 
ument closely resemble that of M. /entigo. The narrow, compressed and upturned 
rostrum of the first five species is unlike the rather slender, dorsally flattened and 
somewhat distally downturned rostrum of M. lentigo; moreover, all of these 
species except M. pilosa have both lateral and gastric spines on the carapace. 
The species with a broad rostrum have hairy or scabrous surfaces quite unlike 
that of M. lentigo. 

Finally, M. lentigo, nitida, pilosa, scabra, and verrilli lack epipods on the 
pereopods whereas the remaining species mentioned above have them on the 
chelipeds. 


Acknowledgments 


We are indebted to K. Smith, Scripps Institution of Oceanography, who as 
chief scientist made cabin space available aboard the R/V Melville, and to ex- 
pedition personnel who helped to collect the material studied. We thank F. A. 
Chace, Jr., and B. B. Collette for critical reading of the manuscript, Keiko Hir- 
atsuka Moore for making the illustrations, and Virginia R. Thomas for entering 
the text on a word processor. 


Literature Cited 


Ambler, J. W. 1980. Species of Munidopsis (Crustacea, Galatheidae) occurring off Oregon and in 
adjacent waters.—Fishery Bulletin 78(1):13—34. 

Benedict, J. E. 1902. Description of a new genus and forty-six species of crustaceans of the family 
Galatheidae, with a list of the known marine species.—Proceedings of the United States Na- 
tional Museum 26(1311):243-334. 

Chace, F. A., Jr. 1942. Reports on the scientific results of the Atlantis expeditions to the West 
Indies, under the joint auspices of the University of Havana and Harvard University. The 
anomuran Crustacea. I. Galatheidae.—Torreia, Havana 11:1—106. 

Faxon, W. 1895. Reports on an exploration off the west coasts of Mexico, Central and South 
America, and off the Galapagos Islands, in charge of Alexander Agassiz by the U.S. Fish 


488 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Commission steamer “Albatross,” during 1891, Lieut.-Commander Z. L. Tanner, U.S.N., 
commanding. XV. The stalk-eyed Crustacea.—Memoirs of the Museum of Comparative Zo- 
ology 18:292 pp., 67 pls. (10 pls. colored). 

Henderson, J. R. 1885. Diagnoses of the new species of Galatheidea collected during the ““Chal- 
lenger’’ Expedition. Annals and Magazine of Natural History (5) 16(96):407-421. 

—. 1888. Report of the Anomura collected by H.M.S. Challenger during the years 1873-76. 
Challenger Expedition, 1872—76.—Report on the Scientific Results of the Voyage of H.M.S. 
Challenger during the years 1873-76. Zoology 27(69):i—xi, 1-221, 21 pls. 

Milne-Edwards, A. 1880. Reports on the results of dredging, under the supervision of Alexander 

Agassiz, in the Gulf of Mexico, and in the Caribbean Sea, 1877, ‘78, ‘79, by the United States 

Coast Survey Steamer “‘Blake,’’ Lieut.-Commander C. D. Sigsbee, U.S.N., and Commander 

J. R. Bartlett, U.S.N., commanding. VIII.—Etudes préliminaires sur les Crustacés.—Bulletin 

of the Museum of Comparative Zoology, at Harvard College 8(1): 1-68, 2 pls. 

, and E. L. Bouvier. 1897. Reports on the results of dredging under the supervision of 

Alexander Agassiz in the Gulf of Mexico (1877-78) in the Caribbean Sea (1878-79) and along 

the Atlantic coast of the United States (1880) by the U.S. Coast Steamer “Blake” .... 

Description des Crustacés de la famille des Galathéidés recueillis pendant |’ expédition.—Mem- 

oirs of the Museum of Comparative Zoology 19(2):1-141, 12 pls. 

Smith, S.I. 1885. On some new or little known decapod Crustacea, from recent Fish Commission 
dredging off the east coast of the United States —Proceedings of the United States National 
Museum 7(32):493—-511. 

Williams, A. B. 1974. A new species of Hypsophrys (Decapoda: Homolidae) from the Straits of 
Florida, with notes on related crabs.—Proceedings of the Biological Society of Washington 
87(42):485—492. 

——. 1976. Integumental organs of unknown function on chelipeds of deep-sea crabs, genus 
Hypsophrys.—Journal of Morphology 150(4):889-899. 

Wood-Mason, J., and A. Alcock. 1891. Natural history notes from H. M. Indian Marine Survey 
Steamer “‘Investigator,’’ Commander R. F. Hoskyn, R.N., commanding.—No. 21. Note on the 
results of the last season’s deep-sea dredging.—Annals and Magazine of Natural History, (6) 
7(27):258-272. 


(ABW) National Marine Fisheries Service Systematics Laboratory, National 
Museum of Natural History, Washington, D.C. 20560; (CLVD) Department of 
Biology, University of California at Los Angeles, Los Angeles, California 90024. 


PROC. BIOL. SOC. WASH. 
96(3), 1983, pp. 489-523 


» THE NEARCTIC SPECIES OF THE BEZZIA BIVITTATA 
GROUP (DIPTERA: CERATOPOGONIDAE) 


Willis W. Wirth and William L. Grogan, Jr. 


Abstract.—The bivittata Group of the genus Bezzia Kieffer, subgenus Bezzia, 
is comprised in North America of at least 15 species, of which the following 12 
are described as new: aklavakensis from Canadian Northern Territory, ander- 
sonorum from Maryland, capitata from Honduras and Arizona, chelistyla from 
Arizona, gibberella from Maryland, /uteiventris from Virginia, mohave from Cal- 
ifornia, nigripes from Utah, sandersoni from Arizona, setosinotum and spathula 
from Maryland, and texensis from Texas. Diagnoses are given of all taxa, a key 
is presented, and all species are illustrated. ) 


This review is one of a series in which additions and corrections are made to 
the “‘Revision of the Nearctic Species of the Genus Bezzia’’ by Dow and Turner 
(1976). In the present series six groups have already been studied in depth since 
1976: Grogan and Wirth (1981) on the genus Amerohelea Grogan and Wirth; Wirth 
and Grogan (1982) on the genus Phaenobezzia Haeselbarth; Wirth (in press) on 
the bicolor Group, Wirth (in press) on the cockerelli and dorsasetula Groups, 
and Wirth eft al. (in press) on the annulipes Group of Bezzia, subgenus Homo- 
bezzia; and Wirth (in press) on the nobilis Group of the subgenus Bezzia s.s. The 
present paper deals with a second group of Bezzia s.s., here named the bivittata 
Group. 

In addition to the revision by Dow and Turner, we urge that users of our Bezzia 
papers also refer closely to the excellent revisions of the Soviet species by Remm 
(1974a, 1974b). We believe that Remm reached a sound basic understanding of 
the characters useful in recognition of natural groups within the genus Bezzia, 
although we would disagree in one or two instances whether certain groups should 
be given generic or subgeneric status. Starting with the characters used by Remm, 
we will offer short diagnoses that should explain the basis of our group classifi- 
cation. We wish to stress that our present treatment is to be considered provi- 
sional and for this reason we will try to be as conservative as possible. 

In addition to Remm’s important papers, we have drawn on several other recent 
works on Bezzia for an evaluation of group characters. Clastrier (1962) presented 
excellent descriptions and figures of a large number of Palaearctic species but 
made very little attempt to place them in systematic groups or to take into account 
the species poorly described by earlier authors. Haeselbarth (1965a, 1965b, 1975) 
treated three groups of African species in similar detail, but added excellent 
diagnoses and taxonomic discussions of the africana and nicator groups, and the 
group which he separated off as the genus Phaenobezzia Haeselbarth. Tokunaga 
(1966) made no attempt at group classification when he offered excellent descrip- 
tions and figures of the New Guinea species. 

Taxonomic characters employed for identification of adult ceratopogonids were 
described by Wirth (1952), Dow and Turner (1976), and Wirth et al. (1977). Wing 


490 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


length is measured from the basal arculus to the wing tip and costal length from 
the basal arculus to the costal apex. Costal ratio is the costal length divided by 
the wing length. Antennal ratio of the female is the sum of the lengths of the 
elongated five distal flagellar segments divided by the sum of the lengths of the 
preceding eight; male antennal ratio is obtained similarly since the five distal 
antennal segments are also elongated in this group. Palpal ratio is the length of 
the third palpal segment divided by its greatest breadth. 

The holotypes and allotypes of our new species are deposited in the U.S. 
National Museum in Washington, D.C. Paratypes as available will be deposited 
in the British Museum (Natural History), London; Muséum National d’ Histoire 
Naturelle, Paris; Canadian National Collection, Agriculture Canada, Ottawa; and 
the California Academy of Sciences, San Francisco. 


Genus Bezzia Kieffer 


Bezzia Kieffer, 1899:69. Type-species, Ceratopogon ornatus Meigen, by original 
designation. 


Diagnosis.—Large, nearly bare, predaceous midges. Body not unusually slen- 
der or dorsoventrally flattened. Antenna short to moderately long. Palpus 5-seg- 
mented, 3rd segment longest, not thickened, with scattered sensilla. Female man- 
dible with 10-20 coarse teeth and with finer proximal teeth in series. Thorax 
robust, mesonotum usually with anterior spine or tubercle, with several strong 
supra-alar and postalar setae. Wing with | radial cell, vein R2+3 absent; costal 
ratio 0.6—1.0; vein M2 originating near r-m crossvein but may be slightly more 
distad or proximad; macrotrichia absent. Legs slender, sometimes with numerous 
spine-like setae; fore femur with 0-12 ventral spines; 4th tarsomere short and 
more or less cordiform; 5th tarsomere without ventral batonnets or strong, sharp- 
tipped setae; female claws simple and equal, usually with small basal tooth on 
inner face; male claws cleft apically. Female abdomen with 1-5 pairs of sclero- 
tized gland rods arising from anterior margins of terga; 2 spermathecae present, 
occasionally a 3rd, rarely only | spermatheca. Male genitalia inverted, 9th tergum 
short with prominent setose cerci, without sclerotized apicolateral processes; 
aedeagus shaped variously, usually triangular in outline; parameres fused to form 
an unpaired distal process, usually rod-like but sometimes variously shaped; dis- 
tistyle well developed and articulated. 

Bezzia is closely related to the large, widespread genus Palpomyia Meigen. 
Most of the Palpomyia species groups can be distinguished from Bezzia by hab- 
itus or genitalic structure; in all cases the presence of vein R2+3 and 2 radial 
cells will distinguish species of Palpomyia and its relatives from Bezzia. Remm’s 
doubts about the independence of Bezzia and Palpomyia arose from his inclusion 
in Bezzia of Phaenobezzia as a subgenus. We support Haeselbarth (1965b) in 
according generic status to Phaenobezzia in view of the remarkable structure of 
the male genitalia with non-articulated dististyle, presence of sharp, spine-like 
setae ventrally on the Sth tarsomere of the female, long costa (costal ratio 0.92— 
0.87), and slender legs without distinct bands or ventral spines on the fore femur. 
Likewise the species Bezzia frontispina Dow and Turner is anomalous in Bezzia 
and we (Grogan and Wirth 1981) placed it in the new genus Amerohelea with a 
Neotropical group of species with | spermatheca, | pair of gland rods placed 


VOLUME 96, NUMBER 3 49] 


extremely laterad on the anterior margin of the 8th tergum, and in most of the 
species with 2 radial cells in the wing. 


Synoptic Key to Subgenera and Groups of Nearctic Bezzia 


1. Male antennal segment 12 no longer than 13, antennal plume weakly de- 
veloped; (mesonotum dull, occasionally weakly shiny, brownish or gray- 
ish with or without vittae; tibiae pale or with a dark medial or basal ring; 
spines of fore femur stout when present; female with 0-5 pairs of gland 
rods; males considerably smaller than females; male aedeagus triangular 
with minute spinules or hairs) (Subgenus Homobezzia Macfie) ........ D, 

— Male antennal segment 12 longest; antennal plume well developed, ex- 
tending at least to apex of 13th segment; (mesonotum black, shiny or dull 
or with silvery hairs, if grayish brown with dark vittae, the hind tibia is 
yellow in midportion, apex broadly black, and all femora bear spines; 
tibiae often black; fore femur with spines slender when present; female 
abdomen with 1-2 pairs of gland rods; males about same size as female; 
male aedeagus variable but not as above) (Subgenus Bezzia Meigen, s.s.) 


Subgenus Homobezzia Macfie 


2. Larger species, female wing |.3—3.4 mm long; mesonotum without bristly 
setae on disc; (fore femur without spines or with 1-4 stout to slender 
spines of similar lengths, with or without strong basal tubercles; pupal 
respiratory horn with numerous (25-60) spiracular openings, apex more 
or less flared, abdominal tubercles well developed) ................... 3 

— Small species, female wing 1.2—1.3 mm long; mesonotum with 2 rows of 
strong bristly setae on disc; (fore femur with 5-7 stout ventral spines of 
alternating uneven lengths arising from distinct elevations; pupal respi- 
ratory horn with only 7—12 spiracular openings, abdominal tubercles small) 

2. 6:0:01 6, 6c 58OkG aI Rael cacti ee ee ae Aare ner gees dorsasetula Group 
SOK MeMuURUInaArMmMeds ventrally, 4.5... aacae emacs es bicolor Group 
— Fore femur armed ventrally with one or more short black spines ...... 4 
4. Fore and mid femora entirely dark brown or with dark bands apical 

5 5.0 Bo io: Beetle CRN op eatin elt i hae cetienada a mnmnna arundel A: An ae cockerelli Group 
— Fore and mid femora with subapical dark bands ......... annulipes Group 


Subgenus Bezzia, s.s. 


5. Fore femur usually unarmed ventrally; legs brown to black; femora and 
tibiae usually with narrow pale rings, rarely femora pale at base or tibiae 
PEA MEME OR clan rey Sac bi sa haat WIE, nein Ba cae aca g eoeivah Rael eke bivittata Group 

— Fore femur armed ventrally with one or more slender black spines; legs 
brown to black, or if banded the pale bands broad ................... 6 

6. Legs broadly yellow or with broad yellow median bands on fore femora 
ZG! TEMAS: See Oe ee ere, Coren amar nobilis Group 

— Legs primarily dark brown to black, at most one pair of legs with broad 
Niello iislaido itn CS Hiaasen ts esi SRG cat lee hdearaen ea, eles cco ccc atiabicti expolita Group 


492 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Dow and Turner (1976) separated the subgenus Pseudobezzia Malloch from 
Bezzia s.s. by the presence of | or more very large stout setae on the vertex, but 
this seta varies so much in size among species that it is only useful but not reliable. 
Most of Dow and Turner’s Bezzia s.s. would fall in Homobezzia Macfie according 
to Remm’s (1974a) characterization. From the structure of the male genitalia and 
female spermathecae Remm’s subgenus Sivabezzia appears to be very similar to 
the expolita Group of Bezzia s.s. Dow and Turner’s subgenus Aspinabezzia is a 
combination of Homobezzia and Bezzia s.s. in which the fore femora lack ventral 
spines. 


Subgenus Bezzia, s.s. 


Diagnosis.—Thorax usually dark brown or black; mesonotum shining or dull 
with silvery hairs or pollinose markings. (In the nobilis Group the mesonotum is 
pollinose grayish brown with dark vittae but the hind tibia is yellow in midportion 
and the apex broadly black, and all the femora bear scattered spines.) Antero- 
marginal spine of mesonotum absent. Males and females more or less of same 
size. Male antennal plume well developed, extending at least to apex of 13th 
segment; 12th segment longest. Fore femur with 0-6 spines; tibiae often black. 
Femaie abdomen usually with a single pair of gland rods. 


Bezzia (Bezzia) bivittata Group 
Figs. 1-10 


Diagnosis.—Small, more or less black species, wing length 1.0—1.4 mm. Vertex 
(Fig. 6) with median seta not very strong. Segment 13 of male antenna with long 
black basal seta (Fig. 3). Legs (Fig. 8) usually black, narrow pale rings usually 
present subapically on some femora and tibiae and subbasally on some tibiae; 
hind femur and tibia usually without pale rings. Fore femur usually without ven- 
tral spines (one rarely present as in Fig. 7). Male aedeagus (Fig. 9) lacking ventral 
setulae and usually with a more or less hyaline, smooth, rounded tip. Distal 
process of parameres and basal arch of aedeagus variously shaped, affording good 
specific characters. 

Species of this group may be separated for the most part from those of the 
expolita Group by the absence of femoral spines, presence of short, distinct necks 
on the spermathecae (Fig. 10), and the short setae on the vertex (Fig. 6). 


Key to the Species of the Bezzia bivittata Group 


1. Halter pale, at least on the knob; female abdomen usually contrasting 
bright yellow, ciehthysternunr brownish). .4- 645 oe eee 2 
= Halterdark brown temale abdomen brown) —. 4.5. eee 6 
2. Femoraand tibiae uniformly yellowish a: = - 240-42] eee 
PL seek A pccentac as rripaat lah ms ich ue cipwai sda flavitibia Dow and Turner (female) 


~ Atleast hind’ femur brown —. 20 ose... oe ye yee eae oo eee 3 
3. Femora solid brown, tibiae uniformly pale; male basistyle bulbous .... 4 
— Legs brown, at most with narrow pale rings; male basistyle various ... 5 
4. Larger species, female wing length 1.23-1.24 mm; female antennal ratio 


1.06—1.10; male aedeagus (Fig. 55) with distal process narrow proxi- 
mally, tip broad and spatula-shaped, basal arms broader .. spathula n. sp. 


VOLUME 96, NUMBER 3 493 


2 Oe EOC 


Se eee OC 


Figs. 1-10. 1-10, Bezzia bivittata; 1-8, 10, female; 9, male: 1-3, antenna; 4, palpus; 5, wing; 6, 
head, anterior view; 7, hind, mid, and fore legs (left to right); 9, 10, genitalia. 


— Smaller species, female wing length 1.05—1.11 mm; female antennal ratio 
0.89; male aedeagus (Fig. 36) slender and rod-like to tip, basal arms 
TNA Tal OW eget 9 aa train acct go oF REAL TATE ET dekh ops dt luteiventris n. sp. 
5. Hind tibia uniformly dark to tip; male basistyle short and stout but with- 
out prominent basal lobe; lobes of aedeagus very long and proximally 


494 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 11. Distribution of Bezzia bivittata. 


slender, hastate distally; parameres with broad capitate distal expansion 

PAU MOOI UNEE, ARGON ES AME dee abet twp aetna A Lect NC ae anOs hh Bd setosinotum Nn. sp. 
— Hind tibia with subapical pale ring; male basistyle with broad basal swell- 

ing with ventral cluster of long setae and shallow distomesal fold; lobes 

of aedeagus short and stout, blunt-tipped; parameres with apex slender 

but with distinct subbasal swelling ................ andersonorum MN. sp. 
6. All femora uniformly dark brown; fore tibia pale brown, mid and hind 

tibiae yellowish; legs with spiny setose vestiture; mesonotum with dense 

fine pubescence and sparse setae ....... flavitibia Dow and Turner (male) 
= Legs andimesonotumothenwise ter oer ce os ck ee eee 7 
7. Tarsi uniformly white, at most fourth and fifth tarsomeres somewhat 

brownish; antenna more or less pale at base; spermathecae elongate 

oval; mesonotum unusually gibbous anteriorly, polished black without 


Pole Mm) :h4022 gp SIR LTS od tea) TS rae ae gibbera (Coquillett) 
— ‘Tarsomeres |I—3 dark at apices, 4—5 uniformly brown; antenna brown at 

base; spermathecae and) mesonotum vanious! —-.. 745-5. 4-4 oa eee 8 
8. Femora and tibiae saturate black, without pale bands ..... nigripes N. sp. 


— Femora with subapical and tibiae with subbasal and subapical pale bands, 
atleast on fore leS: ye. 2 ess a ee eee !) 


VOLUME 96, NUMBER 3 


2? 


IS: 


14. 


. Color saturate black, subapical pale band on fore femur and subbasal 


and subapical bands on fore femur and mid tibia white, definite and 
strongly contrasting; female antennal segments very short ........... 
Color not so black and contrasting; leg bands less distinct or absent; 
female antennal segments Short to lomg <...0....-..08-.4.+ss40.0--- 


. Male parameres with distal swelling or expansion ................... 


Male parameres slender to tip; (female spermathecae short oval with 


CISC ORITE CK) Nas Bera auin. week et eA ee ION Ee TS texensis N. 
. Male paramere with short, broad, spatulate distal expansion; female and 
IMesoOnotalipatternm UNKNOWN 2.2.5.5. 265-.-6-2-22--50 2-6 capitata n. 


Male parameres with bulbous distal swelling; mesonotum with numerous 
pollinose white spots easily seen in anterior view of dry or fresh speci- 
mens; female spermathecae elongate oval, with short necks .......... 


2 pao tub Lon BRUM, pn BME ESSE ST A ie PA an Ce ON ae a sandersoni n. 


Femora and tibiae uniformly dark brown, without pale bands; tarsi uni- 
formly brownish; antennal segments (Fig. 12) short oval, distal five seg- 
ments in female scarcely elongated, antennal ratio 0.79 (spermathecae 
with long tapering necks (Fig. 15); male dististyle short and stout, curved 


tonshanp distalypomt (Figs I6)) 222552. 5)2. 82 22s aklavikensis n. 


Legs with pale rings; antenna with distal segments elongated, female 
te MMe O) OS IPSN: a. Ss Be els, SEs the ee Ne 
Hind tibia with distinct subbasal and subapical pale rings; femora exten- 
sively pale at bases; female with one spermatheca; male aedeagus with 


TUMNCOMC MUNN oem dee, ne et tee athe ee PETE FRO, BRR net mohave n. 


Hind tibia rarely with subbasal and/or subapical pale rings; female with 
two large spermathecae; male aedeagus with tip slender and rounded 

Spermathecae subspherical with short necks; male basistyle broad at 
base, tapering distally; dististyle short, stout nearly to tip; aedeagus with 


495 


sp. 


sp. 


sp. 


sp. 


13 


sp. 


14 


distal process long and tapering with straight sides .. bivittata (Coquillett) 


Spermathecae not as above; male basistyle globular with patch of long, 
mesally directed hairs or stout setae; dististyle hook-like, tapering to 
slender tip; distal process of aedeagus with slender point and concave 
STICIOS sc jaedydeeu Se Puech ke RR a OM RAL rer elie v eh Oe a8 Ran ake ea 


. Mid femur with subapical pale ring; mid tibia with subbasal and subapical 


pale rings; female antenna shorter and stouter; spermathecae with long 
tapering necks; male basistyle with mesal patch of stout setae; dististyle 
short and stout, curved to sharp distal point (eastern U.S.) .......... 


sic tric. pohudtdaca hike UUM Bie Cogs ARNIS em aie Cherie Sab Ae eerie gibberella n. 


Pale rings absent subapically on mid femur and base of mid tibia; female 
antenna elongate; spermathecae elongate oval with short necks; male 
basistyle with mesal patch of long fine hairs; dististyle elongate, slender 


ancdanook-like j(AuiZONA)) \oces doen ceyieta se in ees + 2 a+ chelistyla n. 


Bezzia aklavikensis, new species 
Figs. 12-16 


sp. 


sp. 


Diagnosis.—A large species distinguished from all other species in the bivittata 


group by its unbanded, uniformly dark brown legs, short stout antenna (antennal 
ratio of female 0.79, of male 0.87), and large elongate ovoid spermathecae. 
Allotype Female.—Wing length 1.68 mm; breadth 0.61 mm. Head: Dark brown. 


496 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Figs. 12-21. 12-16, Bezzia aklavikensis; 17—21, andersonorum; 12—15, 18-21, female; 16-17, male: 
12, 21, antenna; 13, 19, palpus; 14, 20, hind, mid, and fore legs (left to right); 15, 18, spermathecae; 
16, 17, genitalia. 


VOLUME 96, NUMBER 3 497 


Eyes broadly separated, a space of four ommatidial facets, a distance of 0.06 
mm. Antennal flagellum (Fig. 12) brown; flagellar segments with lengths in pro- 
portion of 23-11-11-10-10-10-11-11-12-14-15-17-19; antennal ratio 0.79. Palpus (Fig. 
13) brown; lengths of segments in proportion 6-10-16-12-16; palpal ratio 2.67. 
Mandible with 11—12 large coarse teeth. 

Thorax: Dark brown. Mesonotum, scutellum and postscutellum with dense 
short setae and finer pubescence. Legs (Fig. 14) including tarsi uniformly dark 
brown. Wing hyaline, anterior veins brown, posterior veins lighter in color; costal 
ratio 0.68. Halter dark brown. 

Abdomen: Dark brown. Spermathecae (Fig. 15) large, elongate ovoid with short 
necks, measuring 0.104 by 0.063 mm and 0.093 by 0.052 mm. 

Holotype male.—Wing length 1.61 mm; breadth 0.46 mm. Similar to allotype 
female with the following differences: Antennal plume dense, dark brown; fla- 
gellar segments with lengths in proportion of 35-12-12-12-11-12-12-14-20-35-17-16- 
16; antennal ratio 0.87. Palpus with more slender third segment; palpal ratio 3.29. 
Costal ratio of wing 0.62. Genitalia as in Fig. 16. Ninth sternum slightly more 
than twice as broad as long, bulbous in appearance, caudomedian excavation 
deep, broadly V-shaped; ninth tergum tapers abruptly distally to a broad truncate 
margin where it joins the large divergent cerci that extend beyond the basistyles. 
Basistyle 1.6 times longer than broad, broadest basally, tapering slightly distally 
on mesal portion; dististyle about half the length of the basistyle, very short 
truncated in appearance, broadest basally, greatly curved and tapering abruptly 
distally to a broadly pointed tip. Aedeagus triangular, 1.2 times broader than 
long, basal arch 0.3 of total length; basal arm very heavily sclerotized, slightly 
recurved; distal portion more lightly sclerotized, tapering abruptly distally to a 
narrowly rounded, hyaline tip. Parameres heavily sclerotized on proximal por- 
tion, more lightly sclerotized on distal portion; basal arm broad and slightly re- 
curved with a small lateral winglike lobe; distal portion broad proximally, tapering 
distally to broad rodlike form with bulbous rounded tip. 

Distribution.—Canada; known only from the type-locality in the Northwest 
Territory. 

Types.—Holotype male, allotype female, CANADA, NORTHWEST TERRI- 
TORY, Aklavik, 21 June 1953, C. D. Bird (deposited in CNC). 

Discussion.—The specific epithet refers to the type-locality in northwest Can- 
ada where this species was taken. 

The only other species with unbanded dark legs is B. nigripes n. sp. from Utah 
and California. That species differs readily from B. aklavikensis in having darker 
femora and tibiae and paler tarsi, more rounded spermathecae with long, slender 
necks, a more slender palpus (female palpal ratio 5.14), and smaller size (female 
wing length 1.27 mm). 


Bezzia andersonorum, new species 
Figs. 17-21 


Diagnosis. —A medium-sized species most closely resembling B. setosinotum 
in having banded legs and white halter but differing from that species and all 
other species in the group by its short, broad, H-shaped, bipartite aedeagus, 


498 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


parameres with slender distal portion, and female abdomen pale yellow with ovoid 
spermathecae with very short necks. 

Holotype Male.—Wing length 1.15 mm; breadth 0.36 mm. 

Head: Dark brown. Eyes well separated, the width of five ommatidial facets, 
a distance of 0.056 mm. Antennal flagellum uniformly brown; flagellar segments 
with lengths in proportion of 21-10-10-10-10-10-11-13-21-34-13-14-16; antennal ra- 
tio 1.04; plume dense, dark brown. Palpus with lengths of segments in proportion 
of 5-7-13-9-9; palpal ratio 3.25. 

Thorax: Dark brown; mesonotum with moderately dense short setae and finer 
setae giving a pubescent appearance. Legs (Fig. 20) dark brown with banded 
pattern very similar to that of B. setosinotum (Fig. 61) except hind tibia with . 
broad subapical pale band. Wing hyaline, veins pale but easily discernible; costal 
ratio 0.60. Halter white. 

Abdomen: Brown. Genitalia as in Fig. 17. Ninth sternum nearly twice as broad 
as long, caudomedian excavation very deep, U-shaped; ninth tergum tapering 
rather abruptly on basal half, then more gradually on distal half, cerci very short, 
extending to tip of basistyle. Basistyle slightly longer than broad with a broad 
basal swelling with ventral cluster of long setae and a shallow distomesal fold; 
dististyle 0.8 the length of basistyle, broadest in mid-portion, curving and tapering 
on extreme distal portion to broadly pointed tip. Aedeagus very heavily sclero- 
tized, short, broad, H-shaped, and bipartite, slightly longer than broad; basal 
arch to 0.45 of total length, basal arm with tip broadly rounded and recurved 
ventrally; distal portion joined by a slender median bridge that appears to be 
broken due to mounting, each portion tapering slightly distally to narrow rounded 
tip. Parameres very heavily sclerotized; basal arm short, slightly recurved with 
broad, triangular, lateral portion; distal portion constricted at base then broad on 
basal third and tapering abruptly on distal two-thirds to slender rounded tip. 

Allotype female.—Wing length 1.10 mm; breadth 0.43. Similar to holotype male 
with the following differences: Antennal flagellum (Fig. 21) pale on proximal 
portions of flagellomeres, distal portions light brown; flagellar segments with lengths 
in proportion of 14-8-8-8-8-8-8-8-11-11-11-12-15; antennal ratio 0.86. Palpus (Fig. 
19) with lengths of segments in proportion of 5-8-12-7-10; palpal ratio 2.40. Man- 
dible with 10-12 large coarse teeth. Legs dark brown with banded pattern as 
figured, fore femur palest. Costal ratio 0.70. Abdomen pale yellow; two ovoid 
spermathecae (Fig. 18) with very short necks measuring 0.074 by 0.052 and 0.067 
by 0.044 mm. 

Distribution.—Maryland, North Carolina, Quebec. 

Types.—Holotype male, | male paratype, MARYLAND, Worcester Co., Snow 
Hill, 19 June 1968, W. H. Anderson, light trap, from along margin of Nassawango 
Creek one mile upstream from its confluence with the Pocomoke River (Type no. 
76584, USNM). Allotype female, QUEBEC, Rowanton Depot, 28 June 1954, J. 
A. Downes (CNC). Paratype, | female, NORTH CAROLINA, Carteret Co., 19 
June 1977, M. A. Tidwell, light trap. 

Discussion—The species is named in honor of William and Jean Anderson of 
Snow Hill, Maryland, in appreciation of their continued interest in collecting 
Ceratopogonidae for us. The distinctive short, bipartite aedeagus in combination 
with the white halter, yellow female abdomen, and banded legs is sufficient to 
distinguish this species from all others in the bivittata Group. 


VOLUME 96, NUMBER 3 499 


Bezzia bivittata (Coquillett) 
Figs. 1-11 


Ceratopogon bivittatus Coquillett, 1905:60 (female; California). 

Bezzia bivittata (Coquillett):—Kieffer, 1906:58 (combination):—Wirth, 1965:141 
(listed); 1952:238 (male; female redescribed; figs.; California). 

Probezzia bivittata (Coquillett):—Johannsen, 1908:267 (combination):—Malloch, 
1914b:138 (in key); 1915:357 (listed):—Johannsen, 1943:785 (listed). 

Bezzia (Aspinabezzia) bivittata (Coquillett):—Dow and Turner, 1976:126 (rede- 
scribed; status; figs.; distribution). 


Diagnosis.—A medium-sized to large species with dark legs banded as follows: 
pale bands on subapex of fore femur, subbase and subapex of fore and mid tibiae 
(rarely hind tibia banded); spermathecae small, spheroid with short necks, meso- 
notum with two short silvery longitudinal lines; male genitalia with slender ae- 
deagus, short globose basistyles, ninth sternum with shallow caudomedian ex- 
cavation, parameres with slender distal portion and well-developed bifurcate basal 
arms. 

Female.—Wing length 1.40 (1.17—2.11, n = 28) mm; breadth 0.55 (0.78-0.77, 
n = 27 mm. 

Head (Fig. 6): Dark brown including antenna and palpus, vertex silvery polli- 
nose. Antenna (Fig. 1) dark in majority of specimens, occasionally with narrow 
bases of flagellar segments yellowish (Fig. 2); lengths of flagellar segments in 
proportion of 16-9-9-9-9-9-9-9-{2-12-12-14-18; antennal ratio 1.00 (0.77-1.23, n = 
28). Palpus (Fig. 4) with lengths of segments in proportion of 4-6-15-8-12; palpal 
ratio 3.24 (2.40-4.00, n = 28). Mandible with 8—10 large coarse teeth. 

Thorax: Subshining black; anterior fourth of mesonotum with a pair of sub- 
median silvery pollinose spots continued as lines to lateral margins, a pair of 
short, fine, silvery, longitudinal lines continued back from inner edge of spots 
after a short break to half the length of mesonotum; mesonotum and scutellum 
with dense long black pubescence; three black bristles above wing base; scutellum 
with four black marginal bristles. Legs (Fig. 8) dark brown; fore femur with 
subapical, and fore and mid tibiae with subbasal and subapical, narrow yellow 
rings; hind tibia often with faint subbasal and subapical pale rings (Fig. 7); first 
three tarsomeres yellow with apices narrowly dark, last two tarsomeres dark; 
fore femur occasionally with one strong ventral spine (Fig. 7); on hind leg, basi- 
tarsus with two rows of palisade setae, second tarsomere with one row; claws 
(Figs. 7-8) small with basal inner teeth. Wing (Fig. 5) grayish hyaline, anterior 
veins yellowish; costal ratio 0.72 (0.68—0.76, n = 28). Halter dark brown. 

Abdomen: Subshining dark brown; one pair of gland rods as long as four seg- 
ments. Genitalia as in Fig. 10. Spermathecae two (rarely one or three), subspher- 
ical with very short necks, slightly unequal, measuring 0.046 by 0.037 mm and 
0.037 by 0.034 mm. 

Male.—Wing length 0.84—0.124 mm. Similar to female with the following dif- 
ferences: Antennal flagellum (Fig. 3) brown; flagellar segments in proportion of 
28-12-12-12-12-12-14-18-25-55-20-21-24; antennal ratio 1.21; plume dark brown. 
Costal ratio 0.62—0.67. Genitalia as in Fig. 9. Ninth sternum about twice as broad 
as long, hind margin slightly convex with shallow caudomedian excavation. Ba- 
sistyle stout, about as long as basal breadth, base often expanded into a prominent 


500 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


ventromesal lobe; dististyle as long as basistyle, stout, setose, tapering to a distal 
point. Aedeagus about as long as broad with low basal arch; basal arm heavily 
sclerotized, recurved about 60°; distomedian process tapering to moderately slen- 
der tip, sides concave in outline. Parameres heavily sclerotized; basal arm bi- 
lobed; distal portion slender, sides subparallel, tip rounded, hyaline. 

Distribution (Fig. 11).—North America from Alaska to California, east to On- 
tario and Florida. 

Types.—Described from four pinned syntypes, CALIFORNIA, Eureka, **5— 
6,’ H. S. Barber (Type no. 8353, USNM). One pinned syntype has been selected 
and labelled as lectotype. Two syntypes from which the diagnosis was partially 
made have been mounted on slides. 

Specimens examined.—ALASKA: Highway Anchorage to Girdwood, 22 June 
1964, K. M. Sommerman, jeep trap, | male. Fairbanks, June 1967, Sommerman, 
jeep trap, 2 females. 

ARIZONA: Cochise Co., Portal, Southwest Res. Sta., 2—9 June 1972, W. W. 
Wirth, light trap, 4 females. 

CALIFORNIA: Fresno Co., Orosi, 8 July 1947, W. W. Wirth, 1 female (CIS). 
Humboldt Co., Eureka, 5—6, H. S. Barber, 4 female syntypes. Imperial Co., Hot 
Mineral, 30 Apr 1952, J. N. Belkin, 4 males; Westmoreland, 6 Apr 1949, Wirth, 
1 female (CIS). Inyo Co., Saratoga Springs, Death Valley, 30 May 1953, Belkin, 
1 male, 39 females. Mono Co., Fales Hot Springs, 7 June 1948, Wirth, | female 
(CIS); Leavitt Meadow, 7200 ft, 14 Aug 1963, H. B. Leech, flight trap, | female 
(CAS); Topaz Lake, July 1948, R. Coleman, light trap, 1 female; Virginia Creek, 
21 June 1916, H. G. Dyar, 3 females. Monterey Co., Arroyo Seco Ranger Sta., 
1 July 1948, Wirth, light trap, | male, 1 female; Pebble Beach, 26 Aug 1964, R. 
Schoeppner, | male with pupal exuviae. Siskiyou Co., Hornbrook, Aug 1948, R. 
Coleman, light trap, | female (CIS); 4 mi W Weed, 14 May 1948, Wirth, 1 male 
(CIS). Ventura Co., Piru Canyon, 22 Apr 1948, Wirth, | male. 

COLORADO: Rio Grande Co., Beaver Creek, 10,000 ft, 21 June 1972, Wirth, 
Malaise trap, 2 males, 2 females; South Fork, 8000 ft, 20 June 1972, Wirth, 
Malaise trap, 4 males. 

FLORIDA: Alachua Co., Gainesville, Chantilly Acres, 19 Apr, 7 May 1967, 
F. S. Blanton, light trap, 2 females. Liberty Co., Torreya State Park, 27 Apr 
1958, Blanton, light trap, 2 males; 20 May 1966, H. V. Weems, light trap, 4 males, 
1 female; 22 Apr 1967, Wirth, 2 males, 20 females. 

MARYLAND: Prince George’s Co., Patuxent Wildlife Res. Center, May 1976, 
W. L. Grogan, Jr., Malaise trap, 5 males, 6 females; 17 Apr 1977, S. Navai, 
reared from moss on wood over stream, | female; 19 May 1978, Wirth, Malaise 
trap, | female. Worcester Co., Snow Hill, | June 1966, W. H. Anderson, light 
trap, | male, 2 females. 

MICHIGAN: Cheboygan Co., Douglas Lake, 24 June 1954, R. W. Williams, 1 
male. Gogebic Co., 15 June 1960, R. and K. Dreisbach, | female. 

MONTANA: Laurel, 16 July 1917, Dyar, | female. 

NEW BRUNSWICK: Kouchibouguac, 2—13 July 1977, J. R. Vockeroth, 4 males, 
3 females; 11 July 1977, M. Iranochko, | male, 2 females; 9-12 July 1978, J. A. 
Downes, 5 females; 10 July 1978, L. Forster, 5 males, 2 females (CNC). 

NEW MEXICO: Taos Co., Taos, 27 July 1968, Wirth, light trap, 14 males, 7 
females. 


VOLUME 96, NUMBER 3 501 


NEW YORK: St. Lawrence Co., Cranberry Lake, swamp, 25 June 1963, Wirth, 
5 males, 3 females. Tompkins Co., Ellis Hollow, 15 June 1963, C. O. Berg, light 
trap, | female. 

NORTH CAROLINA: Jackson Co., Balsam, 7 July 1968, R. E. Woodruff, 
light trap, | male. 

ONTARIO: Almonte, | July 1954, J. A. Downes, | female (CNC). Ottawa, 
Mer Bleue, 27 May 1960, Wirth, 2 females. 

OREGON: Malheur Co., Little Valley sw Vale, 19 June 1963, K. Goeden, light 
trap, | male, 4 females. 

QUEBEC: Chelsea, 26 May 1960, Wirth, | male. Rowanton Depot, 30 May, | 
June, 7 Aug 1954, J. A. Downes, 2 male, | female (CNC). 

TEXAS: Kerr Co., Kerrville, 21 May 1954, L. J. Bottimer, light trap, | male. 

UTAH: Beaver Co., Beaver, 14 July 1949, G. L. Knowlton, at light, | male. 
Cache Co., Logan, | July 1957, Knowlton, light trap, 1 female. Iron Co., Paro- 
wan, 20 June 1960, Knowlton, | male. Kane Co., Kanab, 21 June 1950, Knowlton, 
1 female. 

VIRGINIA: Alexandria, Dyke Marsh, 13 May 1958, Wirth, 1 female. Fairfax 
Co., Falls Church, 4 July 1950, Wirth, stream margin, | female. Montgomery 
Co., Blacksburg, Apr—June 1960, D. H. Messersmith, | male, 2 females. 

WASHINGTON: Kittataus Co., DeRoux Forest Cpgd., 11 Aug 1971, K. Goe- 
den and A. Gurney, light trap, 2 females. 

WEST VIRGINIA: Pocahontas Co., Cranberry Glades, 16 July 1955, Wirth, 1 
female. 

WISCONSIN: Dane Co., 30 May-7 June 1954, R. J. Dicke, light trap, 5 males, 
8 females; Oregon, 25 May 1962, Wirth, light trap, 1 female. 

WYOMING: Platte Co., Brown Ranch, | Aug 1967, M. Griffith, light trap | 
female. 

Discussion.—As often seen in species with wide distribution, there is consid- 
erable variation in some characters, especially color. The syntypes, and a signif- 
icant number of other specimens from the Pacific Coast states, are considerably 
darker, with the hind tarsus completely brown. A small number of specimens 
were seen with pale bands, usually indistinct, basally and subapically on the hind 
tibia. Occasionally one strong ventral spine occurs on the fore femur; this feature 
at first led to serious consideration as a specific character, but where it occurred 
in about a third of the specimens collected at Torreya State Park in Florida, in 
all instances except one it was found on one leg of a specimen and not on the 
other. Similar presence and absence was seen on occasional individuals from 
other localities. 

A series of males from Kouchibouguac, New Brunswick, however, was rela- 
tively uniform in the correlation of presence of one spine on both fore femora, 
more distinct pale leg bands, more unequal spermathecae, shallower caudal emar- 
gination on the female eighth sternum, and more slender base of the median 
process of the male parameres in one set of specimens; and in another set col- 
lected at the same time and place with a contrasting set of correlated characters 
in the absence of femoral spine, less distinct leg markings, less unequal sper- 
mathecae, deeper caudal emargination on the female eighth sternum, and rather 
stout base of the median process of the male parameres. Further collections and 
study will be necessary to determine if B. bivittata 1s indeed a variable polytypic 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


502 


29 


31 


Figs. 22-31. 22-25, Bezzia chelistyla; 26, capitata; 27-31, flavitibia; 22-24, 28-31, female; 25—27, 
male: 22, 31, antenna; 23, 30, palpus; 25—27, genitalia; 24, 29, spermathecae; 28, hind, mid, and fore 


legs (left to right). 


VOLUME 96, NUMBER 3 503 


species similar to Culicoides variipennis (Coquillett) or if we are dealing with a 
complex of very similar species that we have not learned to distinguish. 


Bezzia capitata, new species 
Fig. 26 


Diagnosis.—A small species with banded legs and dark halter; most closely 
resembling B. setosinotum n. sp. by its parameres with broad capitate tip but 
differing from that species by its aedeagus with broad base and slender pointed 
tip. 

Holotype male.—Wing length 0.86 mm; breadth 0.30 mm. 

Head: Dark brown. Eyes narrowly separated, the wide of three ommatidial 
facets, a distance of 0.03 mm. Antennal flagellum lighter brown on basal half of 
segments 3-7, all of 8 and 9, proximal two-thirds of 10, and extreme bases of 11— 
13; flagellar segments with lengths in proportion of 16-8-8-8-8-8-9-10-12-15-11-11- 
11; antennal ratio 0.90. Palpus lighter brown than rest of head; lengths of segments 
in proportion of 4-6-8-7-7; palpal ratio 1.60. 

Thorax: Dark brown; mesonotum and scutellum with dense short setae and 
fine pubescence. Legs dark brown with banding of femora and tibiae typical of 
B. bivittata (Fig. 8); tarsi paler than those of typical bivittata and more similar 
to those of atypical bivittata (Fig. 7), with only fourth and fifth tarsomeres slightly 
darkened. Wing hyaline, anterior veins light brown, posterior veins paler; costal 
ratio 0.66. Halter brown. 

Abdomen: Brown. Genitalia as in Fig. 26. Ninth sternum twice as broad as 
long, base slightly convex, caudomedian excavation very shallow; ninth tergum 
tapering very abruptly distally to a constriction, then nearly parallel-sided, cerci 
very long and quite broad, extending beyond basistyles. Basistyle slightly curved, 
about 1.3 times longer than broad, very broad basally then tapering and curved 
distally; dististyle nearly as long as basistyle, greatly curved and tapering distally 
to slender pointed tip. Aedeagus heavily sclerotized only on well-defined mid- 
portion; basal arch absent, basal arms nearly straight, directed laterad and ta- 
pering to slender pointed tip; main body triangular in shape with flap-like lateral 
expansions and long slender apical process with narrow pointed tip. Parameres 
heavily sclerotized; basal arm recurved, broad basally, tapering to slender, doubly 
recurved tip; distal portion broad basally, tapering to slender rod which then 
expands to broad, truncate, capitate tip. 

Female.—Unknown. 

Distribution.—Extreme southern Arizona south to Costa Rica. 

Types.—Holotype male, HONDURAS, Comayagua, Rancho Chiquito, 7 June 
1964, F. S. Blanton (Type no. 76585, USNM). Two male paratypes as follows: 
ARIZONA: Santa Cruz Co., Pena Blanca, 10 mi w Nogales, Werner, Nutting 
and Johnson, light trap. COSTA RICA: Puntarenas, Palmar Sur, 5 Aug 1964, F. 
S. Blanton. 

Discussion.—The specific name capitata is in reference to the capitate tip of 
the parameres which is similar to that of B. setosinotum n. sp. The distinctively 
shaped aedeagus of B. capitata is, however, sufficient to distinguish it from se- 
tosinotum as well as all other species in the bivittata Group. 


504 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Bezzia chelistyla, new species 
Figs. 22—25 


Diagnosis. —A medium-sized species with leg markings as in B. bivittata but 
distinguished from that species as follows: Male genitalia wing pincer-like disti- 
styles; aedeagus short, broad with a slender pointed tip; female spermathecae 
large, elongate ovoid with short necks; mesonotum covered with fine pubescence 
only. 

Allotype Female.—Wing length 1.39 mm; breadth 0.48 mm. 

Head: Brown. Eyes narrowly separated, a space of two ommatidial facets, a 
distance of 0.026 mm. Antennal flagellum (Fig. 22) brown, lighter brown on bases 
of all segments; lengths of flagellar segments in proportion of 18-10-10-9-9-10-10- 
10-15-14-15-17-22; antennal ratio 0.97. Palpus (Fig. 23) brown; lengths of segments 
in proportion of 6-8-17-10-11; palpal ratio 4.25. Mandible with 10-12 large coarse 
teeth and 2-3 smaller basal teeth. 

Thorax: Brown; mesonotum covered with fine pubescence only. Legs patterned 
as in B. bivittata (Fig. 8). Wing hyaline, anterior veins brown, posterior veins 
pale; costal ratio 0.69. Halter stem brown; knob dark brown. 

Abdomen: Brown. Spermathecae (Fig. 24) large, unequal, elongate ovoid with 
short necks, measuring 0.070 by 0.052 mm and 0.063 by 0.044 mm. 

Holotype Male.—Wing length 1.57 mm; breadth 0.44 mm. Similar to female 
allotype with the following differences: Antennal flagellum with proximal 7 seg- 
ments indistinctly separated; lengths of flagellar segments in proportion of 27-12- 
12-12-12-13-13-18-30-38-16-19-25; antennal ratio 1.08; plume dark brown. Palpus 
with lengths of segments in proportion of 5-9-19-11-13; palpal ratio 4.75. Wing 
more slender with shorter radial cell; costal ratio 0.64. Genitalia as in Fig. 25. 
Ninth sternum over twice as broad as long, with a shallow caudomedian exca- 
vation; ninth tergum tapering abruptly distally to broad rounded tip where it joins 
the long cerci which extend just below the length of the basistyles. Basistyle 1.3 
times longer than broad, globular, with long ventral and mesal setae; dististyle 
0.83 the length of basistyle, greatly curved, and tapering distally to slender point- 
ed tip. Aedeagus broadly triangular, slightly broader than long, basal arch low, 
about 0.2 of total length, basal arms heavily sclerotized, tips broadly rounded and 
recurved more than 90°; distal portion more lightly sclerotized, tapering distally 
to the long, slender, pointed tip. Parameres heavily sclerotized proximally, more 
lightly sclerotized distally; basal arms with broad lateral lobe and a curved pos- 
terior portion; distal portion slender, rodlike, with slender rounded tip extending 
just beyond basistyles. 

Distribution.—Arizona. 

Types.—Holotype male, allotype female, ARIZONA, Coconino Co., Fort Val- 
ley Exp. Sta., 10 mi NW Flagstaff, 9-12 July 1959, L. A. Carruth, light trap (Type 
no. 76586, USNM). Paratypes, 5 females, ARIZONA: Cochise Co., Portal, 
Southwest Res. Sta., 4 June 1967, C. W. Sabrosky, light trap, 2 females; 5—9 
June 1972, W. W. Wirth, light trap, 2 females; 4 Oct 1967, V. Roth, light trap, | 
female. 

Discussion.—The specific name chelistyla is in reference to the pincer-like or 
claw-like male dististyles. 


VOLUME 96, NUMBER 3 505 


Bezzia flavitibia Dow and Turner 
Figs. 27-31 


Bezzia flavitibia Dow and Turner, 1976:140 (male; New York; figs.). 


Diagnosis.—A medium-sized species distinguished from all other species in the 
group by the following combination of characters: female with pale yellowish 
halter, legs, and abdomen, the head including antenna and thorax contrasting 
dark brown; male with dark brown halter, femora and abdomen, paler tibiae, and 
genitalia essentially the same as those of B. bivittata. 

Female.—Wing length 1.42 (1.35-1.45, n = 7) mm; breadth 0.51 (0.50-0.52, 
n = 7) mm. 

Head: Dark brown. Eyes narrowly separated, a space of three ommatidial 
facets, a distance of 0.04 mm. Antennal flagellum (Fig. 31) brown; flagellar seg- 
ments with lengths in proportion of 19-10-10-9-10-10-10-10-16-15-16-19-23; anten- 
nal ratio 0.99 (0.96—-1.08, n = 7). Palpus (Fig. 30) brown; lengths of segments in 
proportion of 6-10-17-9-10; palpal ratio 3.31 (2.89-4.25, n = 7). Mandible with 8 
large coarse teeth and 2—3 small basal teeth. 

Thorax: Dark brown; mesonotum and scutellum with sparse long setae and 
dense fine pubescence. Legs (Fig. 28) yellowish, femora occasionally light brown- 
ish; tarsi pale on proximal 3—4 tarsomeres, brown on distal 1—2; legs with spiny 
setose vestiture. Wing hyaline, veins pale; costal ratio 0.73 (0.71-0.76, n = 7). 
Halter pale. 

Abdomen: Pale yellowish. Spermathecae (Fig. 29) small, unequal, ovoid to 
spheroid with short necks, measuring 0.059 by 0.044 mm and 0.048 by 0.035 mm. 

Male.—Wing length 1.01—1.57 mm; breadth 0.38—0.45 mm. Similar to female 
with following differences: Antennal flagellum with dark brown plume; lengths 
of flagellar segments in proportion of 23-12-13-13-13-14-15-19-24-47- 18-24-24; an- 
tennal ratio 1.12. Legs with dark brown femora and occasionally tibiae light 
brown. Wing more slender; costal ratio 0.63—0.69. Halter dark brown. Abdomen 
brown. Genitalia as in Fig. 27, similar to those of B. bivittata (Fig. 9). Dow and 
Turner (1976) in their original drawing illustrate a small basal lateral tooth on the 
basal arms of the parameres that is not present in the holotype or any other males 
examined. 

Distribution.—New Brunswick, New York, Ontario. 

Type.—Holotype male, NEW YORK, St. Lawrence Co., Cranberry Lake, 25 
June 1963, W. W. Wirth (Type no. 76587, USNM). 

Specimens examined.—NEW BRUNSWICK: Kouchibouguac, 2-11 July 1978, 
L. Forster, 11 males, 5 females (CNC). 

NEW YORK: Cattaraugus Co., Allegany St. Park, 3 June 1963, W. Wirth, 3 
males (paratypes). Hamilton Co., Newcomb, Hamilton-Essex, 11 May 1959, H. 
A. Jamnback, Berlese trap, | male with pupal exuviae (paratype). St. Lawrence 
Co., same data as holotype, 12 males (6 are paratypes), 7 females. 

ONTARIO: Ottawa, Brittania Bay, 26 May 1960, Wirth, | male; Ottawa, Mer 
Bleue, 27 May 1960, Wirth, 2 males. 

Discussion.—This species is distinguished by the uniformly pale tibiae; in the 
male by the uniformly dark brown femora, especially on the mid and hind legs; 


506 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


on some males the fore tibia is somewhat brownish. The association of the female 
was overlooked by Dow and Turner (1976), and is based on the following: In the 
large collection made by Wirth in the swamp at Cranberry Lake, N.Y., seven 
females were found with leg markings similar to those of male B. flavitibia, but 
with pale halter, yellowish abdomen, and uniformly yellowish legs. There were 
5 males and 2 females of B. bivittata in this collection, but these have distinctly 
dark legs with pale bands on femora and tibiae. The dark brown pigmentation of 
the B. flavitibia thorax is the same for males as for females from Cranberry Lake, 
and the palpi and antennae, allowing for sexual dimorphism, are similar in color 
and shape. Such distinct color dimorphism between sexes is rare but not unheard 
of in Ceratopogonidae. An example is seen in Probezzia pallida Malloch (Tribe 
Sphaeromiini), in which the female is uniformly pale yellowish and the male has 
contrasting black thorax and femora, but is otherwise pale (Wirth and Grogan 
1979). There is also less striking but nevertheless distinct dimorphism between 
the pale female abdomen and the brownish male abdomen of Bezzia nobilis (Win- 
nertz) and B. magnisetula Dow and Turner in the Bezzia nobilis Group. 


Bezzia gibbera (Coquillett) 
Figs. 32-35, 37 


Ceratopogon gibber Coquillett, 1905:60 (female; Cuba). 

Probezzia gibber (Coquillett):—Johannsen, 1908:267 (combination):—Malloch, 
1914b:138 (in table); 1915:357 (noted):—Johannsen, 1943:785 (in list). 

Bezzia gibber (Coquillett):—Kieffer, 1917:330 (combination). 

Bezzia gibbera (Coquillett):—Wirth, 1965:141 (list; distribution); 1974:53 (catalog 
reference).—Dow and Turner, 1976:142 (misidentification in part; female re- 
described; figs.; distribution). 


Diagnosis.—A small species distinguished by its bright yellow antennal scape; 
fourth palpal segment pale, other segments brown; halter stem whitish, knob dark 
brown; setae on scutum arising from raised tubercles; spermathecae large, elon- 
gated, ovoid, subequal; male genitalia with setose lobe on ventromesal base of 
basistyle. 

Female.—Wing length 1.26 (1.11-1.38, n = 9) mm; breadth 0.44 (0.40-0.47, 
n = 9) mm. 

Head: Brown. Eyes narrowly separated, a space of 2.5 ommitidial facets, a 
distance of 0.03 mm. Antennal scape bright yellow, pedicel brown; flagellum (Fig. 
32) yellow on first flagellar segment and proximal '2—*%4 of segments 4-10 and 
basal '4 of distal five segments, distal portions of segments light brown; lengths 
of flagellar segments in proportion of 19-11-11-11-11-11-12-14-17-18-18-19-23; an- 
tennal ratio 0.85 (0.75—0.98, n = 9). Palpus (Fig. 33) brown, fourth segment pale; 
lengths of segments in proportion of 4-7-11-9-8; palpal ratio 2.36 (2.00—2.75, n = 
9). Mandible with 10-12 large coarse teeth and 3-4 smaller basal teeth. 

Thorax: dark brown; mesonotum with setae arising from raised tubercles. Legs 
(Fig. 37) dark brown on hind femur and tibia, lighter brown on mid femur and 
tibia except apex, and proximal % of fore femur and broad midportion of fore 
tibia, yellow on apex of mid tibia, distal fourth of fore femur and base and apex 
of fore tibia; tarsi pale on all tarsomeres; fifth tarsomeres light brown on some 


VOLUME 96, NUMBER 3 507 


Figs. 32-40. 32-35, 37, Bezzia gibbera; 36, 38-40, luteiventris ; 32-34, 37-40, female; 35—36, male: 
32, 40, antenna; 33, 39, palpus; 34, 38, spermathecae; 35-36, genitalia; 37, hind, mid, and fore legs 
(left to right). 


specimens. Wing grayish hyaline, veins brownish; costal ratio 0.72 (0.69-0.76, 
n = 9). Halter stem whitish, knob dark brown. 

Abdomen: Brown. Spermathecae (Fig. 34) large, elongated ovoid with short 
necks; measuring 0.081 by 0.041 mm and 0.074 by 0.037 mm. 

Male.—Wing length 1.15 mm; breadth 0.36 mm. Similar to female with the 
following differences: Antennal flagellum with lengths of flagellar segments in 
proportion of 22-11-11-11-11-11-11-14-20-29-14-15-20, proximal 10 flagellar seg- 
ments indistinctly separated; antennal ratio 0.96; plume golden brown. Fourth 
palpal segment brown. Genitalia as in Fig. 35. Ninth sternum 1.8 times broader 
than long, caudomedian excavation deep, broadly U-shaped; ninth tergum taper- 
ing abruptly distally and becoming rounded where it joins the long, broad cerci 
that extend almost the length of the basistyles. Basistyle curved, base with setose 


508 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


ventromesal lobe; dististyle 0.65 length of basistyle, greatly curved and tapering 
distally to sharply pointed tip. Aedeagus 1.2 times longer than broad; basal arch 
0.20 of total length, basal arms nearly sclerotized, recurved nearly 90°; distal 
portion more lightly sclerotized except along margins, tapering distally to narrow 
rounded tip, margins with lateral flaps that extend almost entire length. Parameres 
heavily sclerotized proximally, more lightly sclerotized distally; basal arms with 
short lateral point and a longer, nearly straight posterior portion; distal portion 
tapering to a slender, rod-like structure with narrow rounded tip. 

Distribution.—Southern Arizona, Texas and Florida south to Panama and 
throughout the islands of the Caribbean. 

Type.—Holotype female, CUBA, Cayamas, 16 Jan, E. A. Schwartz (Type no. 
8355, USNM). 

Specimens examined.—ARIZONA: Maricopa Co., Wickenburg, Hassayampa 
River, 29 June 1953, W. W. Wirth, | male. Pima Co., Quitobaquito, 26 Apr 1959, 
M.S. Adachi, 1 female. Yavapai Co., Oak Creek at Cornville, 10 June 1977, M. 
W. Sanderson, light trap, | female. EL SALVADOR: San Vicente, Santo Do- 
mingo, Oct 1966, F. S. Blanton, light trap, | female. FLORIDA: Monroe Co., 
Big Pine Key, 7 June 1950, St. Bd. Health, light trap, 1 male. JAMAICA: Clar- 
endon Parish, Milk River Bath, 19 Nov 1968, R. E. Woodruff, light trap, | female. 
Runaway Bay, 1-8 Mar 1970, W. W. Wirth, malaise trap, | female. Westmoreland 
Parish, Negril Beach, 10 Dec 1969, E. G. Farnworth, light trap, 1 female. MEX- 
ICO: Oaxaca, Palomares, 5—21 Sept 1961, R. and K. Dreisbach, | female. TEX- 
AS: Kerr Co., Kerrville, May—Sept 1953, 1954 L. J. Bottimer, light trap, 13 
males, 6 females. Llano Co., Enchanted Rock, 1-5 June 1953, W. W. Wirth, 1 
female. TOBAGO: St. John Prov., Charlotteville, Hermitage River Bridge, 12— 
21 Mar 1979, D. Hardy and W. Rowe, Malaise trap, | female. 

Comment.—tThis species is easily recognized by the bright yellow antennal 
scape, the pale fourth palpal segment, the gibbous mesonotum without pruinose 
white spots, but with setae arising from raised tubercles, and the tarsi whitish to 
the tips. Dow and Turner (1976) confused several species under this name. Our 
description is based primarily on the female from Runaway Bay, Jamaica, which 
agreed in external characters with the pinned holotype from Cuba. The descrip- 
tion and figures of the hitherto unknown male are made from the series from 
Kerrville, Texas. 


Bezzia gibberella, new species 
Figs. 41-47 


Diagnosis.—A medium-sized species with dark legs banded on fore leg and 
mid tibia as in B. bivittata but differing from that and all other species in the 
group by having the mid femur with a pale subapical, and hind tibia with a 
subbasal pale band; spermathecae large and ovoid, unequal with long tapering 
necks; male genitalia with triangular aedeagus with hyaline tip, and basistyle with 
dense setae on ventromesal portion. 

Allotype female.—Wing length 1.18 mm; breadth 0.43 mm. 

Head: Dark brown. Eyes narrowly separated, a space of two ommatidial facets, 
a distance of 0.027 mm. Antennal flagellum (Fig. 41) brown, lighter brown on 
basal half of proximal eight flagellar segments and bases of distal five segments; 
flagellar segments with lengths in proportion of 13-9-8-8-8-8-8-9-13-14-15-17-19; 


VOLUME 96, NUMBER 3 509 


MM MAA LEELA AAA 


Sa 


Figs. 41-47. 41-47, Bezzia gibberella; 41-44, 46-47, female; 45, male: 41, antenna; 42, palpus; 
43, wing; 44, spermathecae; 45, genitalia; 46, head, anterior view; 47, hind, mid, and fore legs (left 
to right). 


antennal ratio 1.10. Palpus (Fig. 42) brown, slender; lengths of segments in pro- 
portion of 4-7-13-8-9; palpal ratio 3.25. Mandible with 7-8 large coarse teeth and 
2—3 smaller basal ones. 

Thorax: Dark brown. Mesonotum and scutellum with short dense setae and 
shorter pubescence; postscutellum with dense pubescence, the setae more or less 
in linear groups of 3 or 4. Legs (Fig. 47) dark brown with pale banding on fore 
leg and mid tibia typical of B. bivittata (Fig. 8), mid femur with pale subapical 
and hind tibia with pale subbasal band; tarsi pale on most of proximal three 
tarsomeres, distal two tarsomeres brown. Wing (Fig. 43) hyaline, veins brown; 
costal ratio 0.73. Halter dark brown. 

Abdomen: Dark brown, terga darkest. Spermathecae (Fig. 44) large, ovoid; 
unequal, the larger elongated; with tapering long necks; measuring 0.110 by 0.054 
mm and 0.074 by 0.044 mm. 

Holotype male.—Wing length 1.14 mm; breadth not measurable due to folding. 
Similar to female with following differences: Antennal flagellum more uniformly 


510 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


brown on proximal 8 flagellar segments, segment 9 pale; flagellar segments with 
lengths in proportion of 21-10-10-10-10-10-10-13-19-35-15-18-22; antennal ratio 1.16. 
Palpus with third segment more slender; palpal ratio 4.29. Costal ratio of wing 
0.64. Genitalia as in Fig. 45. Ninth sternum nearly twice as broad as long, cau- 
domedian excavation very deep, quadrately U-shaped; ninth tergum very short 
due to abrupt tapering distally to where it joins the long broad cerci which extend 
much less caudad than the basistyles. Basistyle slightly longer than broad, greatly 
swollen on mesal side basally, then tapering abruptly on mesal side; ventromesal 
area with long setae directed mesad, distomesal area folded with shorter finer 
setae; dististyle 0.67 the length of basistyle, broadest and straightest basally, then 
tapering and greatly curved distally to point tip. Aedeagus triangular, as broad 
as long, basal arch to 0.27 of total length; basal arms very heavily sclerotized, 
recurved more than 90°, tapering to narrow pointed tip; distal portion more lightly 
sclerotized, tapering gradually distally a narrowly rounded, hyaline tip. Parameres 
heavily sclerotized on proximal half, distal half more lightly sclerotized; basal 
arm straight with slightly curved, truncate tip and a small lateral lobe; distal 
portion broad proximally, then tapering distally to a more or less bulbous, round- 
ed tip. 

Variation.—Female wing length 1.30 (1.18—1.41, n = 4) mm; breadth 0.47 (0.43— 
0.53, n = 4) mm. Antennal ratio 1.03 (0.95-1.10, n = 4). Palpal ratio 3.01 (2.89- 
3.25, n = 3). Costal ratio 0.72 (0.71-0.73, n = 6). There is considerable variation 
in the shape of the spermathecae in the type series. Specimens varied from the 
typical spermathecae figured for the allotype to more quadrate spermathecae with 
short, more abrupt necks. 

Distribution.—Maryland to Michigan and Quebec, south to Florida. 

Types.—Holotype male, MARYLAND, Prince George’s Co., Patuxent Wildlife 
Res. Center, 29 July 1978, W. W. Wirth, Malaise trap (Type no. 76588, USNM). 
Allotype female, same data as holotype except taken 30 May 1978; 1 female 
paratopotype taken 8 June 1979; 3 female paratopotypes taken June 1976 by W. 
L. Grogan, Jr. Five male and 3 female paratypes as follows: FLORIDA: Alachua 
Co., Gainesville, Chantilly Acres, 10 May—1 Nov 1967, F. S. Blanton, 2 males, 
| female. MICHIGAN: Cheboygan Co., Douglas Lake, 24, 29 June 1954, R. W. 
Williams, | male, | female. QUEBEC: Rowanton Depot, 6—7 July 1954, J. A. 
Downes, 2 males (CNC). VIRGINIA: Alexandria, 25 May 1952, Wirth, Osmunda 
bog, | female. 

Discussion—The species takes its name from its superficial resemblance to 
Bezzia gibbera (Coquillett). It differs from all other species in the bivittata Group 
in having narrow pale bands on each side of the mid knees, and spermathecae 
with long tapering necks. The male genitalia are distinguished by the conspicuous 
patch of long stout setae on the mesal face of the basistyle, and the unusually 
broad and deep, quadrate excavation on the ninth sternum. 


Bezzia luteiventris, new species 
Figs. 36, 38-40 


Diagnosis.—A small species most closely resembling B. flavitibia Dow and 
Turner in its dark brown femora with contrasting yellow tibiae and pale abdomen 
and halter, but differing from that species as follows: Size smaller, female wing 
1.05-1.11 mm (1.35-1.45 mm for B. flavitibia); tip of female abdomen brown; 


VOLUME 96, NUMBER 3 S11 


spermathecae larger, more quadrately ovoid; antennal flagellum distinctly band- 
ed; male genitalia with deep caudomedian excavation on ninth sternum. 

Holotype female.—Wing length 1.08 mm; breadth 0.39 mm. 

Head: Dark brown. Eyes narrowly separated, a space of three ommatidial 
facets, a distance of 0.037 mm. Antennal flagellum (Fig. 40) pale on proximal 34 
of each of proximal eight segments and proximal 4 of distal five segments; light 
brown on distal portions of all segments giving flagellum a distinctly banded 
appearance; lengths of flagellar segments in proportion of 16-9-9-8-8-8-8-8- 12-13- 
13-14-14; antennal ratio 0.89. Palpus (Fig. 39) light brown; lengths of segments 


and 2-3 smaller basal teeth. 

Thorax: Dark brown; mesonotum and scutellum covered with scattered setae 
and fine pubescence. Legs dark brown on femora; tibiae and proximal four tar- 
someres pale yellow, 5th tarsomeres brown. Wing hyaline, veins pale; costal ratio 
0.70. Halter knob pale yellow. 

Abdomen: Bright yellow; brown on distal three segments, particularly so on 
eighth sternum. Spermathecae (Fig. 38) large; quadrately ovoid, measuring 0.078 
by 0.041 mm and 0.067 by 0.041 mm. 

Allotype male.—Wing length 1.04 mm; breadth 0.30 mm. Similar to female with 
the following differences: Antennal plume brown (distal three segments damaged 
so that flagellar proportions and antennal ratio could not be determined; propor- 
tions in a paratype are 27-10-11-11-11-11-12-15-20-35-16-19-23; antennal ratio 1.05). 
Wing more slender with shorter radial cell; costal ratio 0.61. Abdomen brown. 
Genitalia as in Fig. 36. Ninth sternum 1.7 times broader than long, caudomedian 
excavation very deep, in shape of a broad U; ninth tergum tapering abruptly 
distally on extreme base, then more gradually and becoming rounded where it 
joins the long, slender, divergent cerci, which extend just beyond basistyles. 
Basistyle 1.6 times longer than broad, mesal surface covered with sparse long 
setae; dististyle 0.65 as long as basistyle, curved and gradually tapering distally 
to broadly pointed tip. Aedeagus 1.3 times longer than broad, basal arch shallow, 
0.17 of total length; basal arms very heavily sclerotized, distal portion more lightly 
sclerotized, slender, rod-like with rounded tip. Parameres very heavily sclero- 
tized; basal arms with wing-like lateral process and posterior extension; distal 
portion slightly broader proximally, then more slender and rod-like distally to the 
rounded tip that extends beyond basistyles. 

Distribution.—Virginia. 

Types.—Holotype female, allotype male, 2 male, 2 female paratypes, VIR- 
GINIA, Alexandria, reared from pupae from Osmunda bog, 6-30 June 1951, W. 
W. Wirth (Type no. 76589, USNM). 

Discussion.—The specific epithet is a reference to the yellow abdomen and 
tibiae which serve to distinguish this distinctly marked species from all other 
species in the bivittata Group except B. flavitibia. That species can be distin- 
guished by the characters outlined in the diagnosis. 


Bezzia mohave, new species 
Figs. 48-52 


Diagnosis.—A large species with all tibiae with distinct subbasal and subapical 
bands but differing from all other species in the bivittuta Group in that the female 


512 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Figs. 48-57. 48-52, Bezziu mohave; 53-57, spathula; 48-50, 52-54, 56-57, female; 51, 55, male: 
48, 57, antenna; 49, 54, palpus; 51, 55, genitalia; 52-53, spermathecae; 56, hind, mid, and fore legs 
(left to right). 


possesses Only a single spermatheca and the femora are paler than tibiae distally 
becoming palest proximally; the male resembles the female with genitalia nearly 
identical to those of B. bivittata. 

Holotype female.—Wing length 1.64 mm; breadth 0.57. 

Head: Brown. Eyes narrowly separated, the space of one ommatidial facet, a 
distance of 0.015 mm. Antennal flagellum (Fig. 48) very light brown on proximal 
portions of flagellomeres, slightly darker on distal portions; flagellomeres with 


VOLUME 96, NUMBER 3 513 


lengths in proportion of 21-12-11-11-12-12-12-12-16-16-14-17-21; antennal ratio 0.82. 
Palpus (Fig. 49) brown, slender; lengths of segments in proportion 13-10-18-9-5; 
palpal ratio 3.60. Mandible with 8 large coarse teeth and 4—5 smaller basal teeth. 

Thorax: Dark brown. Mesonotum and scutellum with short dense setae and 
finer pubescence; postscutellum with shorter setae. Legs (Fig. 50) with light brown 
femora proximally becoming slightly darker distally; fore femur with a pale sub- 
apical band; tibiae darker brown with distinct subbasal and subapical light bands; 
tarsomeres 4 and 5 of tarsi pale except darker on extreme apical portions, distal 
3 tarsomeres brown. Wing hyaline, anterior veins light brown, posterior veins 
pale; costal ratio 0.71. Halter brown. 

Abdomen: Brown. A single spheroid spermatheca with short neck measuring 
0.059 by 0.048 mm (Fig. 52). 

Allotype male.—Wing length 1.54 mm; breadth 0.43. Similar to holotype female 
with the following differences: Antennal flagellum with dense, light brown plume; 
flagellomeres with lengths in proportion of 28-13-13-13-13-13-15-19-29-46-18-18- 
24; antennal ratio 1.06. Palpus with more slender third segment; palpal ratio 4.75. 
Genitalia as in Fig. 51. Ninth sternum twice as broad as long, with a shallow 
caudomedian excavation; ninth tergum very short due to abrupt tapering distally 
to a broadly rounded tip where it joins the long, broad cerci which extend slightly 
beyond basistyles. Basistyle 1.4 times longer than broad, broadest basally taper- 
ing distally; dististyle 0.7 the length of basistyle, broadest subbasally, tapering 
and curved on distal half to a slender pointed tip. Aedeagus slightly longer than 
broad, basal arch 0.3 of total length; basal arm heavily sclerotized, recurved 
slightly more than 90°; distal portion lightly sclerotized, tapering slightly distally 
to a rounded tip. Parameres heavily sclerotized on proximal portion, distal portion 
lightly sclerotized; basal arm bilobed as in B. bivittata (Fig. 9); distal portion 
broad basally then becoming slender with a round tip. 

Distribution.—California; known only from the Mojave Desert in Riverside 
County. 

Types.—Holotype female, CALIFORNIA, Riverside Co., Whitewater Canyon, 
6 Apr 1949, W. W. Wirth, at light (Type no. 76598, USNM). Allotype male, 
CALIFORNIA, Riverside Co., Thousand Palms, Willis Palms Oasis, 5 Apr 1955, 
W.R. Richards (CNC). Paratype, 1 female, CALIFORNIA, Riverside Co., Thou- 
sand Plams, 20 Feb 1955, W. R. Richards (CNC). 

Discussion.—The specific epithet is in reference to the Mojave Desert, the only 
known habitat of this species. 

The presence of a single spermatheca easily distinguishes this species from all 
other species in the bivittata Group. with banded legs. 


Bezzia nigripes, new species 
Figs. 64-67 


Diagnosis. —A medium-sized species distinguished by its slender legs with dark, 
unbanded femora and tibiae; spermathecae large, ovoid, subequal, with long slen- 
der necks; palpus slender, third segment very slender (female palpal ratio 5.14). 

Holotype female.—Wing length 1.27 mm; breadth 0.46 mm. 

Head: Dark brown. Eyes broadly separated, a space of about five ommatidial 
facets, a distance of 0.074 mm. Antennal flagellum slightly lighter in shade on 


514 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


extreme bases of distal five segments flagellar segments with lengths in proportion 
of 19-9-9-9-9-9-8-9-] 1-12-13-13-18; antennal ratio 0.83. Palpus (Fig. 66) slender; 
segments with lengths in proportion of 6-10-18-10-13; palpal ratio 5.14. Mandible 
with eight large teeth and three shorter basal ones. 

Thorax: Dark brown. Mesonotum with scattered short setae, but not pubes- 
cent. Legs (Fig. 67) slender with unbanded, dark brown femora and tibiae; tarsi 
dark on apices of tarsomeres | and 2, most of 3 and all of 4 and 5. Wing hyaline, 
anterior veins light brown, posterior veins pale; costal ratio 0.70. Halter dark 
brown. 

Abdomen: Brown, pleura darkest. Genitalia very similar to those of B. vittata 
(Fig. 10). Spermathecae (Fig. 64) large, ovoid with long slender parallel-sided 
necks; subequal, measuring 0.096 by 0.056 mm and 0.078 by 0.052 mm. 

Allotype male.—Wing length 1.35 mm; breadth 0.37 mm. Similar to holotype 
female with the following differences: Antennal flagellum uniformly brown in 
color, plume dense, dark brown; flagellar segments with lengths in proportion of 
28-1 1-11-11-10-11-12-15-26-47-15-15-18; antennal ratio 1.09. Palpus with third seg- 
ment shorter, palpal ratio 3.56. Mesonotum without long setae but with two 
lengths of relatively dense pubescence. Legs with very faint subbasal band on 
fore tibia; hind tarsus entirely brown and mid tarsus more suffused with brown. 
Genitalia as in Fig. 65, very similar to those of B. bivittata (Fig. 9) but differing 
essentially as follows: Aedeagus without well-defined, heavily sclerotized, basal 
portion; parameres broader across basal arms. 

Distribution.—Utah and California. 

Types.—Holotype female, UTAH, Washington Co., Leeds, Red Cliffs Recre- 
ation Area, 22 May 1974, W. L. Grogan, Jr., swept from margin of small stream. 
Allotype male, 2 male paratypes, CALIFORNIA, Imperial Co., Hot Mineral, 30 
Apr 1952, J. N. Belkin (Type no. 76590, USNM). 

Discussion—The specific epithet nigripes, is a reference to the unbanded, dark 
brown femora and tibiae which are sufficient to distinguish this species from all 
other species in the bivittata Group. 


Bezzia sandersoni, new species 
Figs. 68-71, 73 


Diagnosis.—A small species distinguished by its vivid, contrasting pale bands 
on the fore and mid femora and tibiae; female antennal flagellum very short, distal 
segments moniliform to short ovoid (antennal ratio 0.71—0.87); spermathecae large, 
elongate ovoid; male genitalia with deeply emarginate 9th sternum and bulbous 
tip of parameres. 

Allotype female.—Wing length 1.05 mm; breadth 0.47 mm. 

Head: Dark brown. Eyes barely contiguous, joined at midline for the space of 
one antennal facet. Antennal flagellum (Fig. 68) brown, lighter in color on basal 
portions of flagellar segments, darker on apical portions; distal five segments very 
short, ovoid; lengths of flagellar segments in proportion of 15-8-8-8-7-7-7-8-9-9-9- 
10-13; antennal ratio 0.74. Palpus (Fig. 69) light brown, lengths of segments in 
proportion of 4-8-11-8-8; palpal ratio 2.75. Mandible with eight large coarse teeth 
and 5—6 smaller basal teeth. 

Thorax: Dark brown; mesonotum and scutellum covered with short setae and 


VOLUME 96, NUMBER 3 515 


A 


\ 
Hain 


Ze 
yy, 
Vi 


e a ; \\ 
ne atti 
mntenreinn isi 59 


58-63, Bezzia setosinotum; 64-67, nigripes; 58-62, 64, 66-67, female; 71-72, male 


Figs. 58-67. 
58, antenna; 59, wing; 60, 66, palpus; 61—67, hind, mid, and fore legs (left to right), 62, 63, 65, genitalia 


64, spermathecae. 


516 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


fine pubescence. Legs (Fig. 73) dark brown on femora and tibiae, pale bands on 
subapices of fore and mid femora, subbases of all tibiae, and subapices of fore 
and mid tibiae; tars! brown on tarsomeres 4 and 5 and apices of 1-3, pale on 
remainder of 1-3. Wing grayish hyaline, veins light brown; costal ratio 0.69. 
Halter stem light brown, know dark brown. 

Abdomen: Dark brown, pleura purplish. Spermathecae (Fig. 70) large, elongate 
ovoid with short necks; unequal, measuring 0.081 by 0.037 mm and 0.063 by 
0.033 mm. 

Holotype male.—Wing length 0.98 mm; breadth 0.36 mm. Similar to female 
with the following differences: Antennal flagellum pale on basal halves of seg- 
ments 3-10, plume golden brown; lengths of segments in proportion of 19-10-9- 
10-10-11-11-11-16-19-11-12-15; antennal ratio 0.81. Wing more slender with short- 
er radial cell; costal ratio 0.64. Genitalia as in Fig. 71. Ninth sternum twice as 
broad as long, caudomedian excavation deep, broadly U-shaped; ninth tergum 
tapering abruptly and becoming broadly rounded where it joins the long slender 
cerci which extend to the apices of the basistyles. Basistyle bulbous, slightly 
longer than broad, ventral and mesal surface with a few long setae; dististyle as 
long as basistyle, curved and tapering slightly distally to truncate tip. Aedeagus 
triangular, 1.1 times longer than broad, basal arch 0.25 of total length; basal arms 
heavily sclerotized, tapering to recurved, pointed tip; distal portion more lightly 
sclerotized except for margins, tapering at extreme distal end to slender pointed 
tip, apex of which is hyaline. Parameres heavily sclerotized; basal arm with broad 
wing-like lateral extension and rounded posterior portion; distal portion slender, 
rod-like, tip bulbous. 

Variation.—Wing length 1.04 (0.95—1.24, n = 10) mm; breadth 0.46 (0.41-0.56, 
n = 10) mm. Antennal ratio 0.77 (0.71—0.87, n = 10). Palpal ratio 2.36 (2.00-3.25, 
n = 10). Costal ratio 0.69 (0.65—0.71, n = 10). 

Distribution.—Southwestern Utah through Arizona and New Mexico and east 
to Kansas. 

Types.—Holotype male, allotype female, ARIZONA, Coconino Co., Manza- 
nita Forest Camp, 6 July 1977, M. W. Sanderson, light trap (Type no. 76591, 
USNM). Paratypes, 27 males, 28 females, as follows: 

ARIZONA: Cochise Co., Portal, Southwest Res. Sta., May-June 1967, C. W. 
Sabrosky, light trap, 4 males; 5—9 June 1973, W. W. Wirth, light trap, | male. 
Coconino Co., Mormon Lake Village, 26 June 1978, M. W. Sanderson, 3 males, 
3 females; Oak Creek Canyon, 25 June 1959, W. L. Nutting, light trap, | male, 
6 females; 22 July 1959, C. W. O’Brien, light trap, 1 female; Oak Creek, Boot- 
legger Campground, 13 July 1978, Sanderson, 2 males; Oak Creek at Chavez 
Crossing, 20 July 1977, 14 June 1978, Sanderson, | male, 2 females; Oak Creek, 
Encinoso Picnic Ground, 3 Aug 1977, Sanderson; Oak Creek at Grasshopper 
Point, 12 June 1977, 27 Aug 1978, Sanderson, 6 males, 3 females; Oak Creek at 
East Fork, 19 July 1979, Sanderson, 1 male; Manzanita Forest Camp, 6 Aug 1977, 
7 July 1978, Sanderson, 2 males, 3 females. Santa Cruz Co., Ruby, Sycamore 
Canyon, 22 May 1954, G. D. Butler, light trap, 1 female. Yavapai Co., Oak Creek 
at the following points: Baldwin Crossing, Deer Pass Crossing, Josephine Tunnel, 
Oak Creek Village, Page Springs, Red Rock Crossing Verde River, May—Sept 
1977-82, Sanderson, at light, 4 males, 6 females. 

KANSAS: Riley Co., May 1964, N. Marston, Malaise trap, | female. 


VOLUME 96, NUMBER 3 517 


NEW MEXICO: Catron Co., 5 mi E Glenwood, 24 June 1953, W. W. Wirth, 
at light, | male, | female. 

UTAH: Washington Co., Leeds, Red Cliffs Rec. Area, 22 May 1974, W. L. 
Grogan, Jr., near stream, | female. 

Discussion.—This species is named for Milton W. Sanderson in appreciation 
of his interest and cooperation in collecting and sending us extensive biting midge 
collections from Oak Creek Canyon, Arizona. 

Bezzia sandersoni resembles B. gibbera in its large elongate oval spermathecae, 
but differs in its dull mesonotum with numerous pollinose white spots and setae 
not arising from tubercles, its black legs with strongly contrasting pale rings, 
apically dark tarsi, entirely black halter, the extremely short antenna, bulbous 
basistyles without setose basal lobe, and parameres with bulbous tip. 


Bezzia setosinotum, new species 
Figs. 58-63 


Diagnosis.—A medium-sized species with legs banded typically as in B. bi- 
vittata, but differing from that and all other species in the group by the following 
combination of characters: halter white; female antenna short (mean antennal 
ratio 0.86); female abdomen light brown to yellowish with large, elongate-ovoid 
spermathecae with short slender necks; male genitalia with broadly rounded tip 
of parameres and bifurcate aedeagus. 

Allotype female.—Wing length 1.23 mm; breadth 0.47 mm. 

Head: Dark brown. Eyes well separated, a space of about four ommatidial 
facets, a distance of 0.044 mm. Antennal flagellum (Fig. 58) lighter brown than 
head, flagellar segments paler proximally on basal portion, getting progressively 
darker overall distally the distal-most segment darkest; flagellar segments with 
lengths in proportion of 15-10-9-9-9-9-10-9-12-14-13-14-16; antennal ratio 0.86. Pal- 
pus (Fig. 60) light brown, moderately slender; lengths of segments in proportion 
of 5-9-15-8-9; palpal ratio 3.75. Mandible with eight large teeth and 3—4 smaller 
basal teeth. 

Thorax: Dark brown; mesonotum and scutellum with numerous short scattered 
setae; mesonotum not pubescent, but scutellum and postscutellum with fine pu- 
bescence. Legs (Fig. 61) with dark brown femora, and tibiae with banding typical 
of B. bivittata, tarsi pale on proximal three tarsomeres, distal two tarsomeres 
brown. Wing (Fig. 59) hyaline, veins light brown; costal ratio 0.67. Halter white. 

Abdomen: Light brown. Genitalia as in Fig. 62, Seventh sternum lightly scler- 
otized with long setae on distal half. Eighth sternum heavily sclerotized on prox- 
imal third with a shallow caudal notch, covered with long setae. Ninth sternum 
heavily sclerotized, each arm broadly bifurcate. Tenth sternum with five pairs of 
long setae. Spermathecae large, ovoid, elongate, subequal, quadrate with short, 
slender necks, measuring 0.089 by 0.052 mm and 0.070 by 0.048 mm. 

Holotype male.—Wing length 1.24 mm, breadth 0.37 mm. Similar to female 
with the following differences: Antennal flagellum more uniformly dark brown; 
plume dense, dark brown; flagellar segments with lengths in proportion of 29-15- 
11-11-10-11-12-14-21-30-17-18-22; antennal ratio 1.10. Palpal ratio 3.56. Wing more 
slender with shorter radial cell; costal ratio 0.60. Abdomen slightly darker brown. 
Genitalia as in Fig. 63. Ninth sternum 1.7 times broader than long, caudomedian 


518 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


excavation very deep, U-shaped; ninth tergum tapering very abruptly distally into 
conical shape to where it joins the short broad cerci which extend to tip of 
basistyles. Basistyle 1.4 times longer than broad, nearly globose, swollen basally 
on mesal side then tapering rather abruptly on that side, ventral surface covered 
with numerous long setae directed mesally; dististyle 0.7 length of basistyle nearly 
straight, tapering gradually distally to curved, broadly pointed tip. Aedeagus H- 
shaped; basal arms very heavily sclerotized, broadly rounded; distal portion bi- 
furcate and joined by a slender bridge, heavily sclerotized basally, becoming more 
lightly sclerotized distally, a slender apical process arising at bridge and tapering 
distally to narrow tip that extends to ends of basistyles. Parameres heavily scler- 
otized proximally, more lightly sclerotized distally; basal arms broad, hastate in 
shape, becoming constricted at base distal portion broad and bulbous basally, 
then tapering distally before rapidly expanding to the broadly rounded, capitate 
tp. 

Variation.—Female wing length 1.22 (0.95—1.33, n = 10) mm; breadth 0.44 (0.36— 
0.48, n = 10) mm. Antennal ratio 0.86 (0.82—0.98, n = 9). Palpal ratio 2.87 (2.60- 
3.75, n = 9). Costal ratio 0.69 (0.67—0.72, n = 10). There is considerable variation 
in the type-series in the color of the female abdomen, varying from light brown 
to pale yellowish, but in all specimens, even the palest ones, the genitalia are 
well sclerotized. 

Distribution.—New Jersey south to Florida. 

Types.—Holotype male, allotype female, 9 male and 17 female paratopotypes, 
MARYLAND: Prince George’s Co., Patuxent Wildlife Res. Center, June 1976, 
W. L. Grogan, Jr., Malaise trap (Type no. 76592, USNM). Eight male and three 
female paratypes as follows: 

FLORIDA: Alachua Co., Gainesville, Chantilly Acres, 8 May 1967, F. S. Blan- 
ton, | male. 

MARYLAND: Wicomico Co., Salisbury, 20-30 June 1981, Grogan and E. Y. 
Nichols, Malaise trap, 6 males, | female. 

NEW JERSEY: Middlesex Co., 11 July 1958, W. W. Wirth, light trap, | male. 

WEST VIRGINIA: Pocahontas Co., Cranberry Glades 15 July 1955, C. W. 
Sabrosky, | female. 

Discussion.—The specific name setosinotum is in reference to the short, abun- 
dant, stiff, spine-like setae present on the mesonotum and scutellum. It and B. 
andersonorum Nn. sp. are unique among members of the bivittata group in having 
typically banded legs but white halteres. Males of B. andersonorum have a short 
H-shaped but bipartite aedeagus, and the apex of the distal process of the para- 
meres is slender. 


Bezzia spathula, new species 
Figs. 53-57 


Diagnosis.—A small species most closely resembling B. /uteiventris in its dark 
brown femora with contrasting pale yellow tibiae and abdomen, and white halter, 
but differing from that species as follows: male aedeagus with tip broad and 
spatula-shaped and basal arms broader; size larger, female wing length 1.23—1.24 
mm (1.05—1.11 mm for B. luteiventris); female antennal ratio 1.06—1.10 (0.89 for 
B. luteiventris); female antennal flagellum indistinctly banded. 


VOLUME 96, NUMBER 3 519 


Allotype female.—Wing length 1.23 mm; breadth 0.44 mm. 

Head: Dark brown. Eyes narrowly separated, a space of one ommatidial facet, 
a distance of 0.019 mm. Antenna with flagellum (Fig. 57) brown, paler on proximal 
0.75 of proximal segments decreasing to proximal 0.25 on distal segments, thus 
giving flagellum an indistinctly banded appearance; lengths of flagellar segments 
in proportion of 19-10-8-8-9-9-9-9-2 1-16-14-17-18; antennal ratio 1.06. Palpus (Fig. 
54) light brown, base of third segment darker, lengths of segments in proportion 
of 5-9-14-8-10; palpal ratio 2.80. Mandible with 7—9 large coarse teeth and 2-3 
smaller basal teeth. 

Thorax: Dark brown; mesonotum and scutellum covered with numerous scat- 
tered setae and fine pubescence. Legs (Fig. 56) dark brown on femora; tibiae, 
and proximal three tarsomeres pale yellow, distal two tarsomeres light brown. 
Wing hyaline, veins pale; costal ratio 0.71. Halter stem pale, knob white. 

Abdomen: Pale yellow; brown on distal four segments. Spermathecae (Fig. 
53) large, elongate oval; unequal, measuring 0.085 by 0.052 mm and 0.067 by 
0.048 mm. 

Holotype male.—Wing length 1.15 mm; breadth 0.34 mm. Similar to female 
with the following differences: Antenna more brownish on proximal segments; 
plume dark brown; flagellar segments with lengths in proportion of 25-10-11-11- 
1 1-11-13-15-20-29-17-20-20; antennal ratio 0.99. Abdomen brown. Wing more 
slender with shorter radial cell; costal ratio 0.62. Genitalia as in Fig. 55. Ninth 
sternum nearly twice as broad as long, caudomedian excavation very deep, in 
the shape of a truncate U; 9th tergum tapering abruptly distally and becoming 
rounded where it joins the long slender cerci which extend just beyond basistyles. 
Basistyle 1.5 times longer than broad, covered with scattered long setae; dististyle 
0.6 the length of basistyle, curved and gradually tapering distally to bluntly point- 
ed tip. Aedeagus |.2 times longer than broad, basal arch 0.2 of total length; basal 
arms very heavily sclerotized, very broad on proximal portions, then tapering 
and recurving about 60°; distal portion more lightly sclerotized, particularly so 
on the broad, spatula-shaped tip. Parameres very heavily sclerotized; basal arms 
with lateral wing-like expansion and a posterior more truncate extension; distal 
portion bulbous proximally, becoming more slender distally with the rounded tip 
continuing to just beyond basistyles. 

Distribution.—Maryland. 

Types.—Holotype male, allotype female, | female paratype, Maryland, Wi- 
comico Co., Salisbury, 25 May-—7 June 1982, W. L. Grogan, Jr., Malaise trap 
(Type no. 76597, USNM). One male paratype, MARYLAND, Worcester Co., 
Snow Hill, 19 June 1968, W. W. Wirth, swept from margin of Nassawango Creek 
1 mile upstream from its confluence with the Pocomoke River. 

Discussion.—The specific epithet is in reference to the spatula-shaped tip of 
the male aedeagus which serves to distinguish this species from its closest rela- 
tives, B. luteiventris n. sp., and B. flavitibia Dow and Turner. 


Bezzia texensis, new species 
Figs. 72, 74-76 


Diagnosis.—A small species with dark legs banded as in typical B. bivittata, 
but differing from that and all other species in the group by the following com- 


520 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


16 


Figs. 68-76. 67-71, 73, Bezzia sandersoni; 72, 74—76, texensis; 68-70, 73-76, female; 71-72, male: 
68, 76, antenna; 69, 74, palpus; 70, 75, spermathecae; 71-72, genitalia; 73, hind, mid, and fore legs 
(left to right). 


bination of characters: Small size (female wing length 0.95—1.02 mm); short, stout, 
banded antennal flagellum (antennal ratio 0.80); large, unequal, ovoid spermathe- 
cae with long tapering necks; male genitalia with globose basistyles, triangular 
aedeagus with truncate tip, and parameres broad at base with a subbasal, lateral, 
pointed projection. 

Holotype female.—Wing length 0.95 mm; breadth 0.41 mm. 

Head: Brown. Eyes very narrowly separated, a space of about two ommatidial 
facets, a distance of 0.022 mm. Antennal flagellum (Fig. 76) pale on basal halves 
of segments, light brown on distal halves; flagellar segments with lengths in pro- 
portion of 1[3-8-8-8-8-8-8-8-10-10-10-12-13; antennal ratio 0.80. Palpus (Fig. 74) 


VOLUME 96, NUMBER 3 521 


light brown; lengths of segments in proportion of 4-6-9-8-7; palpal ratio 2.25. 
Mandible with 9-10 large coarse teeth and 3-4 smaller basal teeth. 

Thorax: Dark brown; mesonotum and scutellum with dense short stout setae 
on dorsal portions, entire surfaces covered with finer pubescence. Legs dark 
brown, wth banding typically like that of B. bivittata (Fig. 8); tarsi pale on prox- 
imal four tarsomeres, 5th tarsomeres light brown. Wing hyaline, veins pale; costal 
ratio 0.72. Halter brown. 

Abdomen: Brown. Spermathecae (Fig. 75) large, unequal, ovoid with long ta- 
pering necks, measuring 0.081 by 0.048 mm and 0.055 by 0.041 mm. 

Allotype male.—Wing length 0.92 mm; breadth 0.32 mm. Similar to female with 
the following differences: Antennai flagellum with golden brown plume; lengths 
of segments in proportion of 22-10-9-10-10-10-10-13-18-25-13-15-17; antennal ratio 
0.94. Palpal ratio 2.44. Wing more slender with shorter radial cell; costal ratio 
0.63. Genitalia as in Fig. 72. Ninth sternum 1.7 times broader than long, caudo- 
median excavation shallow; ninth tergum tapering abruptly on basal portion until 
it joins the long cerci that extend to tip of basistyles. Basistyle slightly longer 
than broad, globose, ventral and msesal surface with long setae; dististyle about 
the length of basistyle, curved and tapering slightly distally to broad pointed tip. 
Aedeagus triangular, slightly longer than broad; basal arms heavily sclerotized, 
short; distal portion more lightly sclerotized except on margins, tapering distally 
to truncate tip. Parameres heavily sclerotized proximally, more lightly sclerotized 
distally; basal arms broad, recurved about 90° with pointed subbasal lateral pro- 
jection; distal portion broad basally, tapering apically to a slender rod with slightly 
expanded, rounded tip. 

Distribution.—Texas; known only from the type-locality. 

Types.—Holotype female, TEXAS, Gillespie Co., Fredericksburg, 28 July 1967, 
Blanton and Borchers, light trap (Type no. 76593, USNM). Allotype male, | 
female paratype, same data except taken 2 July 1967. 

Discussion.—TYhe small size, pale-banded antennal flagellum, and distinctive 
spermathecae and male genitalia should be sufficient to distinguish this species 
from all others in the bivittata Group with banded legs and dark halteres. 


Acknowledgments 


We are very grateful to J. Antony Downes of the Biosystematics Research 
Institute, Agriculture Canada, Ottawa, and Walter I. Knausenberger of the Col- 
lege of the Virgin Islands Extension Service, St. Croix, for critical reviews of our 
manuscript and their many helpful comments and suggestions. Dr. Downes also 
furnished us with many new Canadian records from the Canadian National Col- 
lection. 

We also extend thanks to the following institutions and persons for the loan of 
specimens: California Academy of Sciences, San Francisco (CAS) by courtesy 
of Paul H. Arnaud, Jr.; California Insect Survey, University of California, Berke- 
ley (CIS) through David Wagner; Cornell University (COR), Ithaca, New York, 
through Q. D. Wheeler; Illinois Natural History Survey, Urbana (INHS) through 
D. W. Webb; Museum of Comparative Zoology, Cambridge, Massachusetts (MCZ) 
through Karen Jepson; University of California, Riverside (UCR) through Saul 
Frommer; and Washington State University, Pullman (WASH) through W. J. 
Turner. 


522 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Literature Cited 


Clastrier, J. 1962. Notes sur les Cératopogonidés XVI.—Espéces du genre Bezzia Kieffer ou ap- 
parentées de la Région Paléarctique. Archives de |’Institut Pasteur d’ Algérie. 40:53—124. 
Coquillett, D. W. 1905. New nematocerous Diptera from North America.—Journal of the New 

York Entomological Society 13:56—69. 

Dow, M. I., and E. C. Turner, Jr. 1976. A revision of the Nearctic species of the genus Bezzia 
(Diptera: Ceratopogonidae).—Virginia Polytechnic Institute Research Division Bulletin 103: 1— 
162. 

Grogan, W. L., Jr., and W. W. Wirth. 1981. A new American genus of predaceous midges related 
to Palpomyia and Bezzia (Diptera: Ceratopogonidae).—Proceedings of the Biological Society 
of Washington 94:1279-130S. 

Haeselbarth, E. 1965a. Notes on Bezzia nicator de Meillon, 1959 (Diptera: Ceratopogonidae). De- 
scriptions of two related new species from Southern Africa.—Novos Taxa Entomologicos 40: 
1-16. 

——. 1965b. Phaenobezzia a new genus of biting midges (Diptera: Ceratopogonidae), with a 
review of the African species.—Zeitschnrift fiir Angewandte Entomologie 52:297-324. 

. 1975. Zur Kenntnis von Bezzia africana und verwandten Gnitzen-Arten aus der Athio- 
pischen Region (Diptera: Ceratopogonidae).—Entomologica Germanica 1:352—370. 

Johannsen, O. A. 1908. New North American Chironomidae.—Bulletin of the New York State 
Museum 124:264—285. 

——. 1943. A generic synopsis of the Ceratopogonidae (Heleidae) of the Americas, a bibliography, 
and a list of the North American species.—Annals of the Entomological Society of America 
36:763-791. 

Kieffer, J. J. 1899. Description d’un nouveau genre et tableau des genres européens de la famille 
des Chironomides (Dipt.).—Bulletin de la Société Entomologique de France 1899:66—70. 

——. 1906. Diptera, Fam. Chironomidae. in P. Wytsman (ed.), Genera Insectorum. Fasc. 42:1— 
78, 4 plates. 

——. 1917. Chironomidés d’Amérique conservés au Musée National Hongrois de Budapest.— 
Annales Historico-Naturales Musei Nationalis Hungarici 14:81—-121. 

Malloch, J. R. 1914. Synopsis of the genus Probezzia, with description of a new species (Diptera).— 
Proceedings of the Biological Society of Washington 27:137—139. 

———. 1915. The Chironomidae, or midges, of Illinois, with particular reference to the species 
occurring in the Illinois River.—Bulletin of the Illinois State Laboratory of Natural History 
10:275—543, 23 plates. 

Remm, H. 1974a. A review of the species of the genus Bezzia Kieffer (Diptera, Ceratopogonidae) 
from the fauna of the USSR. I. (In Russian, English summary).—Entomologicheskoe Obozren- 
ie 53:429-442 (English translation in Entomological Review 53(1):136—145). 

——. 1974b. A review of the species of the genus Bezzia Kieffer (Diptera, Ceratopogonidae) from 
the USSR. II. Subgenus Bezzia s. str. (In Russian, English summary).—Entomologicheskoe 
Obozrenie 53:888—902 (English translation in Entomological Review (53(4): 113-124). 

Tokunaga, M. 1966. Biting midges of the Palpomyiinae from New Guinea (Diptera: Ceratopogon- 
idae).—Pacific Insects 8:101—152. 

Wirth, W. W. 1952. The Heleidae of California.—University of California Publications in Ento- 
mology 9:95—266. 

——. 1965. Family Ceratopogonidae. Pp. 121-142 in A. Stone et al., (eds.), A Catalog of the 
Diptera of America North of Mexico.—United States Department of Agriculture, Agriculture 
Handbook 276: 1696 pp. 

——. 1974. A catalogue of the Diptera of the Americas south of the United States. 14. Cerato- 

pogonidae.—Museu du Zoologia, Universidade de Sao Paulo. 89 pp. 

. InPress. A review of the American predaceous midges of the Rezzia nobilis Group (Diptera: 

Ceratopogonidae).—Proceedings of the Entomological Society of Washington. 

In Press. The North American predaceous midges of the Bezzia bicolor Group (Diptera: 

Ceratopogonidae).—The Florida Entomologist. 

—. In Press. The North American species of the cockerelli and dorsasetula Groups of the 
preedaceous midge genus Bezzia, subgenus Homobezzia (Diptera: Ceratopogonidae).—Pro- 
ceedings of the Entomological Society of Washington. 


VOLUME 96, NUMBER 3 523 


, and W. L. Grogan, Jr. 1979. Natural History of Plummers Island, Maryland. XXIV. Biting 
midges (Diptera: Ceratopogonidae). 2. The species of the tribes Heteromyiini and Sphaero- 
miini.—Proceedings of the Biological Society of Washington 91:847-903. 

, and 1982. The predaceous midges of the genus Phaenobezzia in North America 
(Diptera: Ceratopogonidae).—Memoirs of the Entomological Society of Washington 10:179- 
192. 

, 8. M. Palchick, and L. Forster. In Press. The North American predaceous midges of the 
Bezzia annulipes Group (Diptera: Ceratopogonidae).—Proceedings of the Entomological So- 
ciety of Washington. 

, N. C. Ratanaworabhan, and D. H. Messersmith. 1977. Natural History of Plummers Island, 
Maryland. XXII. Biting midges (Diptera: Ceratopogonidae). |. Introduction and key to gen- 
era.—Proceedings of the Biological Society of Washington 90:615—-647. 


PROC. BIOL. SOC. WASH. 
96(3), 1983, pp. 524-547 


A COMPARATIVE STUDY OF SELECTED SKELETAL 
STRUCTURES IN THE SEASTARS ASTERIAS FORBESI 
(DESOR), A. VULGARIS VERRILL, AND 
A. RUBENS L., WITH A DISCUSSION 
OF POSSIBLE RELATIONSHIPS 


E. K. Worley and David R. Franz 


Abstract.—Morphological structures from the congeneric North Atlantic sea- 
stars, Asterias forbesi (Desor), A. vulgaris Verrill, and A. rubens L., were mea- 
sured and compared. Significant differences in size and/or shape were found 
between A. forbesi and A. vulgaris in the following structures: ventral pedicel- 
lariae, adambulacral spines, oral spines, the madreporite, skeletal ossicles, and 
the shape of the arms. Slight but insignificant differences in size, but not shape 
of these structures was shown between A. vulgaris and the few samples of A. 
rubens examined. 

The firmer, more rounded, less tapering shape of the arms in A. forbesi was 
attributed to the shape of the ossicles and their long processes which form junc- 
tions directly, or by one plate, with adjacent processes throughout the length of 
the arm. In the more flaccid, tapering arms of A. vulgaris (and A. rubens) short, 
blunt ossicle processes connect with adjacent processes by several plates in the 
proximal dorsolateral region forming a more open meshwork and more flaccid 
skeleton. The uniformly small size of these plates throughout the length of the 
arms in young A. vu/garis may account for the less tapered conditions sometimes 
found in small specimens causing them to resemble the shape of the A. forbesi 
arms. Furthermore, variation in size and number of these plates is suggested to 
be associated with the production of morphs in A. vulgaris and A. rubens. The 
forbesi-like animals from the Maine population were therefore diagnosed as local 
morphological variations of the variable species A. vulgaris, and not hybrids. 

Earlier ideas and hypotheses concerning the relationships and origins of the 
three Asterias species are summarized and evaluated; and an hypothesis is for- 
mulated to account for the origin of all Asterias species from a common North 
Pacific ancestor. 


The two seastars, Asterias forbesi and Asterias vulgaris of the Northwestern 
Atlantic coast show many similarities in ecology and morphology. Both species, 
however, exhibit phenotypic variability within and among populations which has 
resulted in problems of identification, especially in those coastal regions where 
the two species are sympatric. While there are no comparative studies to ascertain 
the differences between the species from the extremes of their ranges or from 
the region of overlap on the continental shelf of the Middle Atlantic Bight (MAB), 
the taxonomic traits tabulated by Aldrich (1956) are generally accepted as diag- 
nostic (Schopf and Murphy 1973; Walker 1974) for all areas. 

While isolated and local populations of A. forbesi or A. forbesi-like animals 
may occur in various shallow embayments along the Maine coast (C. Towle, 


VOLUME 96, NUMBER 3 525 


pers. comm.), A. forbesi is distributed more or less continuously from Casco 
Bay, Maine to North Carolina, and may occur south to Florida, although rare 
south of 20°N (Franz et al. 1981). It has been collected over the entire breadth 
of the continental shelf between Cape Hatteras (North Carolina) and Cape Cod 
(Massachusetts), except north of 40°N, where it is confined to the inner shelf, on 
the Nantucket Shoals and Cape Cod Bay. In the SW Gulf of Maine (Isle of Shoals) 
A. forbesi is restricted to depths of 10 m or less (Hulburt 1980), and is generally 
absent from the subtidal zone of the Gulf of Maine. On the shelf south of Cape 
Cod, A. forbesi occurs most frequently at depths <30 m. Asterias vulgaris is 
uncommon or absent from the inshore waters (<20 m) of the Middle Atlantic 
Bight (MAB) but occurs in the deeper, colder waters as far south as Cape Hat- 
teras. Between Cape Cod and Cape Hatteras, both species co-occur in a broad 
zone of overlap on the continental shelf. North of Cape Cod, A. forbesi is rapidly 
replaced by A. vulgaris which occurs commonly on Georges Bank and the con- 
tinental shelf of the Gulf of Maine, northward to the Gulf of St. Lawrence. 

Asterias rubens which is widely distributed along the northeast Atlantic coast, 
resembles A. vulgaris morphologically, hence, they have been considered to be 
closely related species (Coe 1912; Fisher 1930; Nesis 1961) and even identical 
(Clark 1923; Tortonese 1963). In this paper, we examine inter- and intraspecific 
variability in selected skeletal features of all three species in order to evaluate 
the potential usefulness of measurable skeletal characters in taxonomic evalua- 
tion, and to gain insight into the possible relationships and origins of the Atlantic 
species of Asterias. 


Materials and Methods 


Asterias forbesi and A. vulgaris used in this study were collected along the 
northwestern Atlantic Continental Shelf and from the coastal shores of Maine 
and Long Island, New York. Specimens (105) were hand picked from the low 
intertidal shores of Muscongus Bay, Chamberlain, Maine, in August 1978 and 
1979. Other inshore animals (46) were obtained by SCUBA at Shoreham, Long 
Island Sound, in December 1978, and from East Rockaway Inlet (50 animals), 
Far Rockaway, New York (southwestern Long Island) in September 1978 and 
December 1979. Specimens from the northwestern Atlantic Continental Shelf 
were collected, using an otter trawl, by the National Marine Fisheries Service 
during the Ground Fish Survey (Del. II Cruise 78-1, Jan-Feb 1978; Del. II Cruise 
79-01 and Alb. IV Cruise 79-04, Jan—Mar 1979). The Gulf of Maine region (42°02'— 
44°37’) included 25 stations ranging in depth from 35-348 m, and yielded 97 spec- 
imens. Between Cape Cod and Cape Hatteras (42°00’—35°58"), the Middle Atlantic 
Bight (MAB), 1450 specimens of Asterias spp. were obtained from 177 stations 
ranging in depth from 9-220 m. Many of the stations in both the Gulf of Maine 
and the Middle Atlantic Bight yielded only one or two specimens of a species 
and the size range from many stations was very narrow. In order to make valid 
comparisons between the species, measurements of the traits selected for study 
were based on animals within the same size range (R = 4—9 cm). Since the mag- 
nitude of size variability of these characters in a species was found to be no 
greater between stations of a region than between members of the same popu- 
lation, data from all populations of a species from a given geographic area were 
pooled. 


526 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


For comparison, studies were made on formalin-preserved specimens (25) of 
Asterias rubens collected from St. Andrews, Scotland, and a small number of 
museum specimens (alcohol preserved) from Northwest Iceland (10) and south- 
west Greenland (18). 

Observations on living starfish were made only on those collected from Mus- 
congus Bay and Rockaway Beach. Material supplied by the Ground Fish Survey 
in 1979 was frozen immediately on collection and brought to the laboratory in 
this condition. All other specimens were preserved in formalin when collected 
and later studied in the laboratory. 

Four morphological features which are usually considered of diagnostic value 
for Asterias spp. (Coe 1912; Mortensen 1927; Aldrich 1956; Gray et al. 1968) 
were selected for measurement and comparison: the shape of the rays; the size 
and shape of the madreporite; the size and shape of the straight (major) pedicel- 
lariae; and the structure of the skeleton, including the ossicles and the size and 
shape of the adambulacral and oral spines. 

Gross measurements were made with Vernier calipers on the middle arm of 
the trivium. Body radius (R) measured from the center of the disc to the tip of 
the arm, was used for size comparisons between individuals of the same and 
different populations and species. Tapering of the rays was calculated from the 
ratio (a/b) between the diameter of the arm at the base (a) and the diameter 
measured | cm from the tip (b). Thus, more tapered rays exhibit higher a/b ratios 
than blunt, parallel-sided rays. 

Spines, straight pedicellariae, and the madreporite were removed from at least 
10 animals from each region and measured with a calibrated Wild Dissecting 
Scope. Spines and pedicellariae were selected at random from the proximal third 
of the arms, and each recorded value, based on the measurement of the calcareous 
skeleton, represents the average of a minimum of 10 samples per animal. 

After removal of podia and internal organs, animals were skeletonized by plac- 
ing One Or more arms or the entire specimen, depending on size, in undiluted 
commercial Clorox (sodium hypochlorite) for 10-25 minutes, or until the soft 
outer covering was dissolved. Skeletons were washed in several changes of water 
and dried at room temperature. 

Color variations were noted and compared in live and frozen specimens only, 
but no detailed studies were made. Preserved material showed color loss and 
could not be considered reliable for comparisons. 


Observations 


Shape of arms.—The a/b ratios, measured on animals over a size range of R = 
2-8 cm, from MAB populations of A. forbesi and A. vulgaris were weakly cor- 
related with body size (R), but the degree of tapering was greater in all sizes of 
A. vulgaris (Table 1). The a/b ratios from Shoreham and Rockaway Beach pop- 
ulations of A. forbesi were not significantly correlated with body size and had 
similar mean a/b ratios (Shoreham x = 2.23; Rockaway Beach x = 2.16). Asterias 
vulgaris from Muscongus Bay likewise showed no statistically significant corre- 
lation between a/b and R, and had essentially the same a/b mean value (« = 2.13) 
as that of the two shallow, coastal water populations of A. forbesi. In shape, 
therefore, small A. vulgaris from Muscongus Bay resembled A. forbesi of com- 
parable size. 


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528 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Asterias rubens, in the range of R = 2—9 cm, from the St. Andrews and West 
Greenland populations exhibited statistically significant positive correlation be- 
tween R and a/b ratios. The slopes for these populations were distinctly steeper 
than for MAB A. vulgaris of comparable size (Table 1). 

Madreporite.—Madreporites from a wide range of animal sizes were measured, 
but only data from specimens of R = 5—9 cm were used for comparison of the 
diameter (size) and height (convexity) between the species. Few A. forbesi from 
the Shoreham and Rockaway Beach populations were under 4 cm and none of 
the A. vulgaris from Muscongus Bay or available specimens of A. rubens were 
over 9 cm. Within this size range, there was little intrapopulation variability in 
either size or convexity of the madreporites in any species. 

The average diameter of the madreporites from animals of comparable size 
from the three populations of A. forbesi studied was x = 4.13 mm, which was 
greater than diameters from populations of A. vulgaris (€ = 3.30 mm) or A. ru- 
bens (* = 2.82 mm), a variation hardly perceptible to the naked eye (Table 2). 

The height (convexity) of the madreporites showed no significant correlation 
with body size (R) within any populations. Asterias forbesi from all regions had 
slightly higher (more convex) madreporites (¢ = 1.18 mm) than A. vulgaris (x = 
0.76 mm) and A. rubens (x = 0.80 mm) from all populations (Table 2). 

Pedicellariae.—Dorsal: Numerous dorsal, straight (major) pedicellariae of the 
short, rounded type (Coe 1912) were distributed over the abactinal surface be- 
tween the dorsal spines in A. forbesi from all populations. These pedicellariae 
ranged in size from 0.33—0.49 mm (Table 2), with little size difference between 
the population. 

Narrow, pointed major pedicellariae (Coe 1912) were the usual dorsal type in 
both A. vulgaris and A. rubens, but both broad and pointed forms were sometimes 
found together. In both species, major pedicellariae were always less numerous 
and more widely distributed than in A. forbesi. Dorsal pedicellariae from animals 
of comparable size from populations of A. vulgaris from the Gulf of Maine (0.47— 
0.63 mm) and the MAB (0.58—0.72 mm), and from the samples of A. rubens from 
Northwest Iceland (0.44—0.88 mm) were longer than those of A. forbesi (Table 
2). On the other hand, the range of length of the dorsal pedicellariae (0.29-0.41 
mm) in the Muscongus Bay population of A. vulgaris was shorter than in A. 
forbesi, but comparable to the range found in the St. Andrews specimens (0.31— 
0.47 mm) of A. rubens (Table 2). There was no correlation between length of the 
dorsal pedicellariae and body size (R) in any population (Table 3). 

Ventral: All ventral, straight pedicellariae were longer than the dorsal pedicel- 
lariae. However, in all three species, we selected for study only the major ped- 
icellariae from both the inner and outer adambulacral spines of the proximal 
region of the arm. Each spine usually carried from one to eight major pedicellariae 
on the outer surface of the distal half. The number and size of the pedicellariae 
gradually decreased distally in the arm. 

In A. forbesi, the major pedicellariae were broad and blunt. Comparing the 
pedicellariae from animals of comparable size, the differences in mean length 
were Statistically insignificant between those from subtidal populations (Rocka- 
way Beach x = 0.54 mm; Shoreham x = 0.50 mm) and those from deeper (15-60 
m) MAB stations (¢ = 49 mm) (Table 2). In no population was there a significant 
correlation between body radius (R) and length of the pedicellariae (Table 3). 


529 


VOLUME 96, NUMBER 3 


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VOLUME 96, NUMBER 3 


532 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


The major ventral pedicellariae in A. vulgaris and A. rubens were longer and 
more pointed than those of A. forbesi (Table 2). Variation in the mean length of 
the pedicellariae did not differ significantly between populations of A. vulgaris 
(Gulf of Maine x = 0.68 mm; MAB x = 0.67 mm; Muscongus Bay x = 0.59 mm) 
and A. rubens (St. Andrews « = 0.60 mm; N.W. Iceland x = 0.77 mm) (Table 2). 
A weak but significant correlation with body radius (R) was noted in all popu- 
lations of A. vulgaris, but only in the St. Andrews population of A. rubens (Ta- 
ble 3). 

Skeleton and spines.—The main divisions of the Asterias skeleton (ambulacral, 
actinal, marginal, dorsolateral, carinal) are based on the principal longitudinal 
rows of inbricated ossicles, rigidly held together throughout life, except in the 
dorsolateral region where there is a loose, irregular connection between ossicles 
which forms an open meshwork. Details of ossicle morphology have not been 
adequately described in Asterias and a standard terminology has not been estab- 
lished. Consequently, many of the terms used in this study to describe skeletal 
structures follow those presented by Turner and Dearborn (1972) for the mud- 
star Ctenodiscus crispatus, in addition to those employed for A. amurensis (Fish- 
er, 1930) and for A. forbesi and A. vulgaris by Hyman (1955). 

Ossicle designates the large, calcarious structures which make up the basic 
rigid framework of the skeleton (Fig. 1A). Ossicles are typically quadrilateral, 
except in the dorsolateral region where 2—6 sided forms are found. They regularly 
have one or more pustules, mound-like elevations or bosses, with a central 
depression, the spine pit, for the attachment of the spine (Fig. 1A). A projection, 
or process from each side of the ossicle forms a suture or junction with a process 
from an adjacent ossicle either directly, by overlapping or underlapping, or in- 
directly by one or more narrow, bar-like, overlapping plates. These are flattened, 
calcareous structures, smaller than ossicles and usually without processes, pus- 
tules, or spines (Fig. 1B). Connections between the ossicles are called arches 
(Hyman 1955). Longitudinal rows of parallel arches were designated channels by 
Fisher (1930). In the spaces between the arches, the fenestrae (Hyman 1955), a 
membrane stretched between the walls of the arches, is perforated by a regularly 
arranged ring of a fairly specific number of openings for the papulae (Fig. 1B). 

Actinolateral ossicles.—In all three species of Asterias, the actinolaterals usu- 
ally do not overlap but form a row along the outer edge of the adambulacral 
ossicles (Figs. 2-4). Each actinolateral forms sutures internally with two or three 
underlying adambulacral ossicles. The actinolaterals are small, dorsoventrally 
thickened and have short, blunt anterior and posterior processes. In A. forbesi 
the dorsal (abactinal) process of each ossicle is elongated and forms a suture 
directly with the elongated ventral (actinal) process of the adjacent inferomarginal 
ossicle to form a small, round arch, within which the fenestra membrane bears 
a single ring of 4-6 openings (Fig. 1B). This row of arches forms the actinal 
channel of Fisher (1930). 

In A. vulgaris, and A. rubens, the dorsal processes of the actinolateral ossicles 
are short and connect with the short, blunt ventral processes of adjacent in- 
feromarginal ossicles by single plates, each of which may bear a spine in the prox- 
imal part of the arm (Figs. 3, 4). The arches thus formed are dorsoventrally 
oblong, larger than those in A. forbesi, and the membrane within the fenestra has 
a ring of 5-8 openings. 


VOLUME 96, NUMBER 3 533 


Fig. |. Asterias forbesi. A, Diagrammatic views of a typical dorsolateral ossicle: outerside (above 
left), inner side (above right), side view (center). B, Diagram of a typical arch: 1, spine pit; 2, pustula; 
3, process; 4, articulation depression; 5, ossicle; 6, plate; 7, membrane in fenestra; 8, opening for 
papula. 


Inferomarginal ossicles.—TYhe inferomarginal ossicles in all three species of 
Asterias form a prominent longitudinal row of large, closely imbricated, quadri- 
lateral ossicles dorsal to the actinolaterals on the ventral (actinal) side of the 
animal. Each ossicle bears 3—4 pustules with slit-shaped, obliquely arranged spine 
pits, the most posterior of which lies nearest the ambulacral groove (Figs. 2-4). 
In A. forbesi, the dorsal (abactinal) processes of the interomarginal ossicles are 
elongated and pointed. Each articulates either directly, or by a single plate, with 
the ventral (actinal) process of the adjacent superomarginal ossicle (Fig. 2), thus 
forming a row of rounded arches, designated the intermarginal channel by Fisher 


534 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 2. Asterias forbesi. Semidiagrammatic drawing of a section of skeleton from proximal region 
of left side of animal (MAB). a, ambulacral ossicles; b, adambulacral ossicles; c, actinolateral ossicles; 
d, inferomarginal ossicles; e, superomarginal ossicles; f, dorsolateral ossicles and plates; g, carinal 
ridge. x3. 


(1930). The membrane within each fenestra contains an elongated ring of 8-10 
openings. 

In A. vulgaris and A. rubens, the dorsal processes of the inferomarginal ossicles 
are short and blunt and connect with the ventral processes of the superomarginals 
by at least one plate, thus forming oval arches which are larger than those of A. 
forbesi, especially in large animals (Figs. 3, 4). The membrane within each fe- 
nestra may have as many as 16—18 openings arranged in an elongated ring. 

Superomarginal ossicles. —The superomarginal ossicles in all three species of 
Asterias form a strong, prominent longitudinal overlapping row along each margin 
(ambitus) of the ray dorsal to the inferomarginals (Figs. 2-4). Each ossicle has a 


VOLUME 96, NUMBER 3 5)3)5) 


Fig. 3. Asterias vulgaris. Semidiagrammatic drawing of a section of skeleton from proximal region 
of left side of animal (MAB). a—g. See Fig. 2. x3. 


short anterior process which overlaps a longer posterior process of the preceding 
ossicle, a ventral (actinal) process which forms a suture with a dorsal (abactinal) 
process forming a junction, either directly or indirectly by one or more plates 
with an actinal process of the adjacent dorsolateral ossicle. 

In A. forbesi, the superomarginal ossicles are smaller and flatter than the in- 
feromarginals. Each ossicle has 3—5 round spine pits, one on the anterior process, 


536 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


A 


Fig. 4. Asterias rubens. Semidiagrammatic drawing of a section of skeleton from proximal region 
of left side of animal (St. Andrews). a—g. See Fig. 2. x3. 


one or two on the central boss, and one on each lateral process. Both of the 
lateral processes are elongated and articulate directly, or by a single short plate, 
with adjacent processes, thus forming two rows of arches. Each fenestra mem- 
brane contains 16-20 openings which are scattered throughout the entire area. 

The anterior and posterior processes of the superomarginals in A. vulgaris and 
A. rubens are elongated, but the lateral processes are short and blunt (Figs. 3, 
4). Each ossicle has three round spine pits, two on the central boss and one on 
the actinal process. Articulation with the adjacent dosolateral ossicles is by 1-3 
elongated plates forming a row of large arches. The 16—20 openings in each fe- 
nestra membrane are arranged in a ring along the periphery of the arch. 


VOLUME 96, NUMBER 3 537 


Dorsolateral ossicles.—TYhe dosolaterals in all three species of Asterias are 
typically irregular, rectangular (also 2-6 sided), non-overlapping ossicles whose 
processes articulate with processes from adjacent ossicles, either directly or by 
one or more plates, forming an irregular meshwork which is not symmetrical on 
the two sides of the arm. In smaller (2-3 cm) animals, ossicles show more regular 
anterior-posterior orientation to form 2—3 longitudinal rows on either side of the 
mid-dorsal carinals. In larger specimens, however, this arrangement is usually 
obscured by unequal lengthening of processes and/or plates. Each ossicle has at 
least one central pustule with a round spine pit, and frequently, one or two small 
spine pits are on the processes (Figs. 1A, 2-4). 

In A. forbesi, the processes of the ossicles are elongated, tapered, and articulate 
directly, or by not more than one plate, with processes of adjacent ossicles to 
form small, firm, round arches. The membrane in each fenestra has 8-10 open- 
ings. 

The processes of the dorsolaterals in A. vulgaris and A. rubens are shorter, 
more rounded and usually articulate with processes of adjacent ossicles by 1-3 
plates, thus forming larger, more oblong and irregular arches which are quite long 
in large specimens. The 16-20 openings in the fenestra membrane are arranged 
in a ring around the periphery. In large specimens, additional openings may be 
present within the center of the ring. 

Rarely, a skeleton was obtained from widely separated populations (Shoreham, 
A. forbesi; St. Andrews, A. rubens) which showed no discernible pattern in the 
arrangement of the ossicles of the abactinal side. They were small, nearly square 
with short, rounded processes, and no intervening plates between the processes. 
The arches were small and irregular, and the ossicles often were superimposed 
on other ossicles. These individuals had a very rigid and compact skeleton. 

Carinal ossicles.—A median row of anterior-posteriorly aligned, overlapping 
carinal ossicles extends between the dorsolaterals from the edge of the disc to 
the tip of each ray, forming a central, dorsal ridge or keel (carina! ridge) in all 
three species of Asterias (Figs. 2-4). In A. forbesi, the lateral processes are 
elongated and pointed, forming sutures directly or by a single plate, with pro- 
cesses of adjacent dorsolaterals. A regular row of arches (channel) on either side 
of the carinal ossicles is usually evident in the skeleton. Each carinal ossicle has 
3-4 pustules each with a round spine pit, arranged in a regular pattern; one on 
the anterior process, one or two directly posterior on the central body of the 
ossicle, and at least one small spine pit on each lateral process. 

In A. vulgaris and A. rubens, the lateral processes of the carinal ossicles are 
short, blunt, and connect with adjacent dorsolateral processes by 2—3 overlapping 
plates forming a distinct row of arches on either side of the carinal ossicles. Each 
ossicle has one or two pustules, one in front of the other along the median ridge 
of the ossicle. This arrangement results in a more or less straight, single row of 
spines along the median dorsal ridge of the arm (the carinal ridge). 

Adambulacral spines. —TYhe adambulacral spines in the proximal region of the 
arms in all three species of Asterias were more or less regularly arranged in the 
typical Asterias alternating 1-2-1-2 etc. pattern (Mortensen 1929) forming three 
rows, an inner series along the edge of the ambulacral groove, a middle, and an 
outer row. Only spines from the inner and outer rows were measured and com- 
pared (Fig. 5A, B). 


538 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


(bs A. FORBES|! A. VULGARIS A. RUBENS 
CF" | 
“| qi D, b, mI ai 
B 
cS, di d, qd, d, d. oF d.. 


Fig. 5. Diagrammatic outer views of typical adambulacral and oral spines from Asterias forbesi, 
A. vulgaris and A. rubens. A, Adambulacral spines: a,, typical form of outer spines; a2, grooved 
form; b,, typical form of inner adambulacral spines; b,, pointed form. B, Oral spines: c,, typical outer 
oral spine; c,, pointed form; d,, typical inner oral spine; d,, pointed form; d3, curved form. Scale line 
equals | mm. 


The length of the inner and outer adambulacral spines in A. forbesi was com- 
parable in the three populations studied (Table 2), ranging between a low of 1.58 
mm to a high of 3.33 mm. In all populations, both inner and outer ambulacral 
spines were truncate, wide, (¢ = 0.66 mm) and flattened. The outer spines were 
frequently grooved on the outer side (Fig. 5). Thinner, more pointed spines were 
sometimes found among the more truncate forms. 

In A. vulgaris and A. rubens, both inner and outer adambulacral spines were 
typically round, pointed (Fig. 5), and approximately within the same length range 
(1.33-2.93 mm) as those in A. forbesi in animals within the same size range (Table 
2). The slightly narrower average width (« = 0.5 mm) of the A. vulgaris spines 
tended to give the rows a more delicate and crowded appearance, but in animals 
in the same size range, there was no significant difference among the species in 
the actual number of spines present per row. Double-pointed as well as flat, but 
ungrooved, truncate spines sometimes occurred along with the pointed spines. 

The adambulacral spines in all species exhibited a strong correlation with body 
radius (R) (Table 3). While the slope for both types of spines was steep in all 
populations (Table 3), the Muscongus Bay population (A. vulgaris) showed the 
steepest slope for both inner and outer adambulacral spines (Table 3). 


VOLUME 96, NUMBER 3 539 


Oral spines.—The inner and outer oral spines in A. forbesi were about the 
same length or slightly longer than the adambulacral spines (Table 2), and resem- 
bled them in shape, but were not grooved. Occasionally an animal was found 
with larger and more pointed oral spines. Significant correlation between length 
of the outer oral spines and body radius (R) was evident in the Rockaway Beach 
(r = 0.772) and Shoreham (r = 0.913) populations (Table 3) but not in those from 
the MAB (r = 0.256). Inner oral spines showed significant size correlation only in 
the Shoreham population (r = 0.818) (Table 3). There was no significant difference 
in the three populations in the mean ratio between length of inner/outer oral spines 
(Rockaway Beach x = 0.72 mm; Shoreham x = 0.76 mm; MAB x = 0.74 mm). 

In both A. vulgaris and A. rubens, however, there was a difference in size and 
shape between the inner and outer oral spines (Table 2). The inner oral spines 
were shorter, slightly curved, narrow, and sharply pointed while the outer oral 
spines were longer, straight, narrow, round, and pointed (Fig. 5C, D). Neither 
truncate nor grooved oral spines were found in any population. The correlation 
of body radius (R) and oral spine length was significant in all populations (Table 
3). The mean inner/outer spine ratios in A. vulgaris (Gulf of Maine x = 0.58; 
Muscongus Bay « = 0.54; MAB « = 0.516) were lower than those in A. forbesi, 
but not significantly different from A. rubens (St. Andrews x = 0.46, West Green- 
land x = 0.53; Northwest Iceland x = 0.54). There was greater difference in length 
between the inner and outer oral spines in both A. vulgaris and A. rubens than 
was found in A. forbesi. 

Color.—Color patterns and intensities in both A. forbesi and A. vulgaris showed 
wide variation between different stations but in each species, a distinct intrapopu- 
lation color pattern tended to predominate in a given locality (Coe 1912). In A. 
vulgaris from the MAB, the basic color of the abactinal surface was yellowish to 
reddish brown with varying amounts of bluish purple ranging from a narrow band 
along the carinal ridge to being suffused over the aboral surface from the dark 
purple disc to the tips of the arms, hence the name of ‘‘purple starfish.’’ Speci- 
mens of A. vulgaris from deep locations (200 m) from the Gulf of Maine were 
cream colored to light tan. Animals from Muscongus Bay ranged from light brown 
to deep reddish brown and purplish blue. Spines on animals from all locations 
were light yellow and not conspicuous against the pale color of the dermis. 

Asterias forbesi from the MAB were reddish and abactinal markings were darker 
blue than those of the lighter, yellowish colored, sympatric A. vulgaris. Live 
Rockaway Beach animals were usually a uniform, deep red, but occasionally 
lighter forms were found. Light (white or yellowish) colored spines outlined the 
arms and frequently formed a definite carinal ridge. Irregular longitudinal rows 
of dorsolateral spines often made a distinct pattern against the dark color of the 
aboral surface. 


Discussion 


Characters that showed significant size-relationship to body radius (R) includ- 
ed inner and outer adambulacral spines and inner and outer oral spines in all 
three species; and the tapering of the arm (a/b ratio) in A. vulgaris (except from 
Muscongus Bay), and A. rubens. 

Characters that did not show significant size relationships in any species in- 
cluded dorsal pedicellariae; ventral pedicellariae; madreporites. 


540 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Characters that were significantly different in size and/or shape among the 
species included ventral pedicellariae; inner oral spines; inner and outer adam- 
bulacral spines (shape); ossicles (shape); madreporite (shape). 

Some of the structures which show correlation between size and body radius 
(R) are associated with functions which increase with growth of the animal. The 
adambulacral spines, for example, extend over the ambulacral canal and protect 
the underlying podia (Hyman 1955). As the canal widens with growth of the 
animal, the spines lengthen to continue coverage of the canal. The oral spines, 
which cover the oral region, also lengthen with increase in body size to maintain 
their protective function. Tapering of the arms (greater a/b ratio), on the other 
hand, in A. vulgaris and A. rubens increases as the animal grows due to the 
lengthening of the greater number of connections (plates) between the ossicles in 
the proximal region of the arms than in the distal portion. In A. forbesi where 
growth of the arm is more uniform throughout its length, there is less tapering (a 
lower a/b ratio). 

Ventral and dorsal pedicellariae and the madreporite, which are not signifi- 
cantly size-related, serve essential functions which do not change as the animal 
grows. These structures attain their optimal size at an early age of the animal and 
do not change significantly. 

Structures (ventral pedicellariae, madreporite) which show significant differ- 
ences among the three species in size and/or shape have long been considered 
diagnostic. Coe (1912) suggested that the difference in size and shape of the major 
ventral pedicellariae in A. forbesi and A. vulgaris was the most reliable criterion 
for species identification. Comparison of measurements of these structures from 
widely separated populations justifies this conclusion. While the shape of the 
ventral pedicellariae is similar in A. vulgaris and A. rubens, there are differences 
in size. Ventral pedicellariae from A. vulgaris of Muscongus Bay have a slightly 
lower size range than (other) A. vulgaris from the Gulf of Maine or MAB. However, 
this range falls within that of ventral pedicellariae from the St. Andrews popu- 
lation of A. rubens, but is shorter than those of A. rubens from N.W. Iceland. 
Variations in size would therefore appear to be population related and not species 
specific. 

The difference in the color of the madreporites in A. forbesi and A. vulgaris 
has usually been considered more significant for species identification than shape 
and size (Coe 1912; Aldrich 1956; Gray et al. 1968). However, the greater con- 
vexity of the structure in A. forbesi was described in detail by Verrill (1866). This 
difference in shape is slight but significant and may not be readily detected except 
by comparison of measurements. 

Some structures which are significantly different in A. forbesi and A. vulgaris 
are variable and have not been listed as diagnostic. In this group are included the 
outer and inner adambulacral and outer and inner oral spines. Verrill (1866) and 
later Clark (1904) noted the predominance of blunt, grooved adambulacral spines 
in A. forbesi compared to the slender, often pointed spines of A. vulgaris. In the 
specimens available for the present study, the shape of most of the adambulacral 
spines of a specimen corresponded to this distinction, although both types of 
spines were often found together in the same animal. This situation is not an 
indication of hybridization, but rather the expression of a polymorphic structure 
(Schopf and Murphy 1975). 


VOLUME 96, NUMBER 3 541 


Clark (1904) noted that the oral spines in A. forbesi and A. vulgaris were ‘‘not 
peculiar.’ However, in the animals of the present study, the inner oral spines 
showed variability both in size and shape. In A. vulgaris and A. rubens, the inner 
oral spines were on the average shorter than the outer oral spines and usually 
curved, while in A. forbesi, the inner orals were straight and approximately equal 
in length to the outer orals. The difference between the lengths of the inner orals 
measured in A. vulgaris and in the few representatives of A. rubens available 
suggest that it would be interesting to evaluate the mean size of these structures 
between and within larger samples of both A. vulgaris and A. rubens populations. 
When inner oral spines of A. vulgaris from all areas were compared with those 
of the St. Andrews population only, the difference between the mean size was 
slightly greater in A. vulgaris (¥ = 1.85 mm) than in the A. rubens population 
(x = 1.33 mm) and statistically significant at the P = 1.0 level but not at the P = 
0.5 level. 

If the observations of Fisher (1930) on A. amurensis, the North Pacific species, 
are compared with the present results, the resemblance, especially in the adam- 
bulacral spines and the inner oral spines, to the North Atlantic species is evident. 
Fisher described the adambulacral spines of A. amurensis as long, compressed, 
tapered, bluntly pointed and grooved, a description resembling that for the adam- 
bulacral spines usually found in A. forbesi. The inner oral spines were described 
as curved, tapering and blunt, the shape which is comparable to that regularly 
found in A. vulgaris and A. rubens and infrequently in A. forbesi. These, and 
other close resemblances suggest a common ancestor for the species. 

Descriptions in the literature of the skeletal structures of Asterias spp. are 
generalized and incomplete. Mortensen (1927) called the dorsal skeleton of A. 
rubens faintly developed. Verrill (1866) distinguished the skeleton of A. forbesi 
from that of A. vulgaris by the larger number of plates in the dorsal area which 
gave it a “‘stout’’ condition. In A. vulgaris, the lateral plates were described as 
separated by large spaces and were connected by plates broken into distinct 
pieces. Hyman (1955) recognized a basic, reticulate pattern but the diagram she 
presented was reproduced from Fisher (1928) and was based on Pacific forms of 
Asterias. Gray et al. (1968) called the skeleton of A. forbesi a mosaic of inter- 
locking plates of ossicles, and that of A. vulgaris a network of narrow, bar-like 
plates forming a weak skeleton. 

Basically, the skeletons of the three species of Asterias are equally strong and 
well developed, and the arrangement of the main ossicles all follow the same 
general pattern (Figs. 2-4). The essential differences in the skeletal structure 
among the species are the shape of the lateral processes (long, pointed in A. 
forbesi; short, blunt in A. vulgaris and A. rubens), and the greater number of 
plates between the processes in A. vulgaris and A. rubens. These differences are 
associated with the rigidity, shape, and tapering (a/b ratio) of the arms. In A. 
vulgaris and A. rubens, where the plates elongate during the growth of the animal, 
lateral distance between the ossicles, especially in the dorsolateral region, in- 
creases and forms the open meshwork with elongate fenestrae resulting in a more 
flaccid skeleton. This less rigid arrangement also gives the arms a flatter and 
wider appearance, especially in the proximal portion. However, in the distal part 
of the arms, the plates between the processes remain small or absent forming a 
tight, rigid meshwork and producing the tapering of the arms and an increased 


542 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


a/b ratio. In A. forbesi, on the other hand, where the skeletal meshwork is formed 
by sutures between elongated processes with only one or no connecting plates, 
a more rounded and rigid framework results throughout the length of the arm 
giving a lower a/b ratio. Elongation of the several plates between processes in A. 
vulgaris and A. rubens may be one explanation for these species attaining a 
greater size than is possible in A. forbesi where growth of the skeleton is limited 
principally to elongation of the processes directly connecting the ossicles. 

Local populations of A. rubens with rigid, straight and rounded arms have been 
described from several locations (N.W. Iceland, Heding 1892 museum label; 
Scandinavia, Masden, pers. comm.; Great Britain, Vevers 1947). These morphs 
may result from variations in the size and/or number of the connecting plates. 
Similarily, the forbesi-like animals from the Muscongus Bay population may be 
juveniles of A. vulgaris in which the plates in the proximal dorsolateral region 
have not elongated. No specimens of otherwise typical A. forbesi have been 
described as having flaccid skeletons. Hence, the invariably rigid skeleton would 
appear to have become a genetically fixed characteristic in A. forbesi. 

The more frequent occurrence of a prominent carinal ridge in A. vulgaris and 
A. rubens than in A. forbesi may also be related to the greater dependence of 
the more flexible skeletons on a stronger, more rigid central keep for support. 

Our observations, as well as those of all other workers, clearly demonstrate 
distinctive differences between Asterias forbesi and A. vulgaris, differences both 
at the morphological and structural as well as the physiological and ecological 
levels. These differences are clearcut in spite of marked genetic similarity (Schopf 
and Murphy 1975). The co-occurrence of such closely-related congeners in the 
shallow shelf of the N.W. Atlantic is rather remarkable, particularly in view of the 
overlapping ecological and spatial niches of the species, and the probable high 
degree of competition in areas of sympatry, and the generally low overall diversity 
of asteroids in the Middle-Atlantic continental shelf of the NW Atlantic. We 
attempt to provide a historical hypothesis to account for the co-occurrence of 
these species in the following section. 

The specific identity of A. vulgaris vis-a-vis A. rubens is not resolved in this 
study; moreover, descriptive analyses of morphological characteristics probably 
can not produce a definitive answer. Our results indicate that populations of A. 
vulgaris from the Gulf of Maine and MAB differ from the St. Andrews population 
of A. rubens in skeletal structures such as the dorsal and ventral pedicellaria 
(Table 2), but that these small differences disappear in significance when the 
Muscongus Bay population of A. vulgaris and the N.W. Iceland population of A. 
rubens are included in the comparison. Thus, from a strict descriptive taxonomic 
viewpoint, we can provide no basis for maintaining A. vulgaris as distinct from 
A. rubens, even though populations of A. vulgaris are geographically isolated 
from A. rubens. Even assuming the questionable existence of a permanent pop- 
ulation of A. rubens in S.W. Greenland, the directional flow of surface currents 
along the W. Greenland coast in the Davis Strait, and the absence of Asterias 
populations in Labrador, imply essentially complete geographical isolation of 
populations of A. vulgaris in the Gulf of St. Lawrence from the nearest major 
concentrations of A. rubens in Iceland. 

Since the major difference between the geographical zone of A. rubens in the 
N.E. Atlantic and A. vulgaris in the N.W. Atlantic is the thermal environment 


VOLUME 96, NUMBER 3 543 


rather than the structure of biotic communities, we would predict that genetic differ- 
entiation would involve primarily physiological rather than morphological traits— 
and these have not been investigated. Until this is done, we believe that the most 
parsimonious approach to the question of taxonomy would be to conserve the 
status quo, 1.e., while recognizing the very close genetic and taxonomic relation- 
ships between A. vulgaris and A. rubens, to continue to consider them as separate 
species until more relevant aspects of their biology can be evaluated. 


Hybridization Between Asterias forbesi and Asterias vulgaris 


Hybridization between A. forbesi and A. vulgaris is often assumed to occur, 
and to account for the existence of individuals with external characters inter- 
mediate between typical A. forbesi and A. vulgaris. Given the genetic similarity 
demonstrated by Schopf and Murphy (1975), such hybridization might be ex- 
pected. However, evidence for hybridization is very weak; and, to the best of 
our knowledge, is limited to the occurrence of morphological ‘‘intermediates.”’ 
Even the work of Ernst (1967), cited as containing experimental evidence for 
hybridization, contains no conclusive evidence proving that hybridization occurs 
under natural conditions. 

After examining thousands of individuals of both species from the Middle At- 
lantic continental shelf, including many from the geographical zone of sympatry, 
we have never observed an individual which could not be assigned with confi- 
dence to one or the other species. While it is possible that hybrids may occur in 
shallow waters near Cape Cod, this was discounted by Clark (1923:235), who 
stated unequivocally that **. . . if such hybrids occur they must be very rare, for 
there are no authentic specimens on record or extant, as far as I know.”’ More- 
over, sympatric populations of A. forbesi and A. vulgaris which occur in the SW 
Gulf of Maine (Isle of Shoals area) are distinguishable morphologically and eco- 
logically (Hulbert 1980, pers. comm.). 

The possibility that A. forbesi-like animals occurring in embayments along the 
Maine Coast may represent hybrids seems unlikely to us. Inshore populations of 
typical A. forbesi occur in the SW Gulf of Maine and south of Cape Cod, but not 
on Georges Bank or the inner shelf of the Gulf of Maine, where summer bottom 
temperatures remain below 15°C (Franz et al. 1981). More likely, these unique 
coastal populations of A. forbesi-like seastars are either morphological variants 
of A. vulgaris (as is the case in the Muscongus Bay intertidal population reported 
in this paper) or relict populations of A. forbesi which are retained within and 
restricted to shallow coastal embayments. Relict populations of oysters (Cras- 
sostrea virginica) occur in the Sheepscot Estuary, and McAlice (1981) has pre- 
sented evidence for the existence of relict populations of other estuarine trans- 
hatteran invertebrates in Maine coastal estuaries. If relict populations of A. forbesi 
date from the Hypsithermal Period (7000-9000 years BP), it would not be sur- 
prising if they had undergone a degree of morphological and genetic differentiation 
from the main body of A. forbesi populations further south. 


The Origin of Asterias forbesi and Asterias vulgaris 


Several workers have speculated on the origins of Asterias forbesi and A. 
vulgaris. A. H. Clark (1923) observed that no species of Asterias ranged farther 


544 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


south than A. forbesi, indicating to him the likelihood that A. forbesi evolved 
from A. vulgaris as a general consequence of adapting to warmer waters. No 
mechanism of speciation was suggested. 

More recently, Schopf and Murphy (1973) postulated that A. vulgaris evolved 
from A. forbesi during the Pleistocene as a result of geographic isolation brought 
about by the emergence of Georges Bank during the last glacial episode. This 
land barrier effectively isolated northern populations from the remaining southern 
populations. Presumably, natural selection favoring individuals living in the cold 
and increasingly harsh environment brought about the evolution of Asterias vul- 
garis. With the submergence of this land barrier during the Holocene, the recently 
differentiated ““semispecies’’ again converged to produce the partially overlap- 
ping species which presently exist. 

While this hypothesis does provide for allopatric speciation and accounts for 
the major thermal adaptations of the species, as well as their close genetic rela- 
tionship, there are several problems. The time period allocated for speciation 
may be too short—roughly 7000 years (the period between the minimum sea level, 
ca 15,000 years BP, to about 7000 years BP when rising sea levels would have 
again united the separated areas). Another, more serious, objection is presented 
by the severity of environmental conditions believed to have existed on the coast- 
al shelf north of 42°N during glacial periods. While many questions remain, recent 
studies (McIntyre 1976) indicate that essentially arctic conditions prevailed during 
glacial maxima. Asterias vulgaris presently does not live in arctic waters. It 
reaches its northern limit in the Gulf of St. Lawrence, and is not found in the 
Strait of Belle Isle (Grainger 1966). This suggests that A. vu/garis could not have 
persisted north of 42°N, i.e., north of the land barrier, during glacial maxima. 
Schopf and Murphy’s hypothesis also fails to account for, or explain the rela- 
tionships between the NW Atlantic species and the North Pacific and NE Atlantic 
congeners A. amurensis and A. rubens. 

Tortonese (1963) suggested that Asterias rubens (including, in his opinion, A. 
vulgaris), A. forbesi and A. amurensis comprise a superspecies, i.e., a set of 
allopatric species sharing a common ancestor which, in his opinion, was probably 
A. rubens. He further suggested that the center of dispersion was the North 
Atlantic, and that A. forbesi and A. amurensis differentiated from A. rubens (or 
a closely-related ancestor) following westward dispersion into the NW Atlantic 
(A. forbesi) and, either eastward dispersion across Siberia or westward through 
the Canadian Arctic into the North Pacific (A. amurensis). He provided no ex- 
planation as to how or when such dispersions may have occurred or under what 
conditions A. forbesi differentiated from sympatric A. vulgaris. 

While agreeing with Tortonese that the species of Asterias comprise a super- 
species (sensu Mayr 1963) we do not agree with his suggestion of an Atlantic 
origin. While an Atlantic origin of some boreal North Pacific invertebrate species 
is probable (Durham and MacNeil 1967), there is an emerging consensus that the 
amphiboreal fauna of the North Atlantic is largely derived via transarctic dis- 
persals in the Pliocene (or, in some cases, earlier) (Durham and MacNeil 1967; 
Franz et al. 1980; Franz et al. 1981). 

We suggest that the common ancestor of all North Pacific and North Atlantic 
Asterias probably lived in the North Pacific during the Miocene. After the opening 


VOLUME 96, NUMBER 3 545 


of the Bering Straits at the end of the Pliocene, the climate warmed and this 
species, or one of its descendants, migrated from the North Pacific into the North 
Atlantic via the Arctic, entering the North Atlantic via the straits and sounds of 
the Canadian Archipelago and/or Norwegian Sea (Nesis 1961). 

As populations spread southward along both coasts of the North Atlantic, 
geographic and genetic connections were broken, possibly facilitated in the late 
Pliocene, by the closure of the Isthmus of Panama and the development of the 
Labrador Current system. These events brought about a strengthening and mod- 
ification of the axis of the Gulf Stream (Berggren and Hollister 1977) and provide 
an explanation for environmental changes in the Atlantic which, as argued by 
Franz and Merrill (1980), may account for the evolution of a separate boreal fauna 
in the North Atlantic, as well as the ecological separation of NW Atlantic and 
NE Atlantic Asterias populations to produce A. forbesi and A. rubens. 

During one or more of the extensive interglacial periods, or possibly as late as 
the Holocene, A. rubens may have extended its range westward along the island 
arc of the North Atlantic (Faroes, Iceland, Greenland) and via the Davis Strait 
to Labrador and New England, becoming partly sympatric with A. forbesi. During 
glacial maxima, A. forbesi would have been displaced southward (as would have 
A. rubens on the European coast). Possibly, relict populations of A. rubens re- 
maining in the NW Atlantic may have been able to survive these periods in the 
NW Atlantic by extending their ranges southward into the Middle Atlantic Bight. 
Alternately, the westward range extensions of A. rubens into the NW Atlantic 
may have occurred in the Holocene. In either case, these relict populations of 
A. rubens in the NW Atlantic are now known as A. vulgaris. 

Evidence to support this hypothesis is circumstantial. The diversity of asteroids 
in the North Pacific is very great, leading to the conclusion that this area marks 
the center of origin of the Asteridae. Since species of Asterias do not occur in 
subtropical or tropical waters, it follows that North Atlantic populations were 
derived from North Pacific ancestors via transarctic migration. Caenozoic migra- 
tions are thought to have occurred at least twice: the well-documented late Plio- 
cene/early Pleistocene Beringian Transgression, | million years BP (Allison 1978); 
and a less well documented late Miocene transgression which is substantiated 
primarily on zoogeographic evidence (MacNeil 1965; Durham and MacNeil 1967). 

Evidence for the westward range extensions of boreal invertebrates during the 
Pleistocene is summarized in Franz and Merrill (1980). In spite of the North 
Atlantic Drift, some European species have extended their ranges westward via 
Iceland to Greenland (Kraeuter 1974). Populations of A. rubens presently occur 
abundantly in Iceland and, at least periodically, in West Greenland (Einarsson 
1948). Since A. rubens could not have survived in these areas during the Wis- 
consin glacial maximum, it follows that A. rubens extended its range westward 
during the Holocene. It seems reasonable to assume that if A. rubens can extend 
its range westward as far as West Greenland during present conditions, it may 
have been able to extend its range into the Davis Strait and to Labrador, and 
hence southward to New England, during the unusually warm conditions which 
prevailed during the Hypsithermal. Alternately, if the westward range extension 
occurred earlier, e.g., during a late interglacial, A. rubens may have been able 
to survive glacial conditions in the Middle Atlantic zone of the NW Atlantic, 


546 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


based on the analysis of environmental conditions in this zone during the Wis- 
consin glacial maximum (McIntyre 1976). We have no basis for distinguishing 
between these alternatives. 

Unfortunately, we are unable, at this point, to propose a definitive procedure 
to falsify the hypotheses discussed above. Ultimately, when taxonomic relation- 
ships within the A. amurensis complex are more clearly established, it may be 
possible to investigate evolutionary relationships among geographical groups of 
Asterias using both biochemical (isoenzyme) and morphological (cladistic) ap- 
proaches. . 


Acknowledgments 


We thank Dr. Arthur E. Merrill of the National Marine Fisheries Service (ret.) 
for supplying the starfish from the Groundfish Surveys (1978 and 1979), and P. 
J. Fallon, Jr., for samples of A. forbesi from Shoreham, New York. We also 
appreciate the kindness of Dr. Chris Todd, University of St. Andrews, Scotland, 
and Dr. F. Jensensium Madsen of the Zoologisk Museum, University of Den- 
mark, Copenhagen, for providing specimens of A. rubens. This research was 
supported in part by a grant from the City University of New York (PSC-CUNY 
Grant Nos. 13315, 14059). 


Literature Cited 


Aldrich, F. R. 1956. A comparative study of the identification characteristics of A. forbesi and A. 
vulgaris.—Notulae, Natural Academy of Natural Sciences of Philadelphia 285: 1-3. 

Allison, R. C. 1978. Late Oligocene through Pleistocene molluscan fauna in the Gulf of Alaska 
region.—The Veliger 21(9):171-188. 

Berggren, W. A., and C. O. Hollister. 1977. Plate tectonics and paleocirculation—a condition in 
the ocean.—Tectonophysics 38:11—48. 

Clark, H. L. 1904. Echinoderms of the Woods Hole region.—Bulletin of the United States Fish 
and Wildlife Service 1902 22:545-5S76. 

—. 1923. The distribution and derivation of some New England echinoderms.—American Nat- 
uralist 57:229-237. 

Coe, W.R. 1912. Echinoderms of Connecticut.—Connecticut State Geological and Natural History 
Survey, Bulletin 19, 152 pp. 

Durham, J. W., and F. S. MacNeil. 1967. Coenozoic migrations of marine invertebrates through 
the Bering Strait Region, in The Bering Land Bridge (ed. David M. Hopkins), Stanford Uni- 
versity Press, pp. 327-349. 

Einarsson, E. 1948. Echinoderms.—Zoology of Iceland 4(70): 1-67. 

Ernst, E. J. 1948. The distribution, ecology, environmental behavior and possible hybridization of 
the sea stars Asterias forbesi (Desor) and Asterias vulgaris Verrill in the subtidal zone of Long 
Island.—Ph.D. Dissertation, New York University (Univ. Microfilms 468-6060, 84 pp.). 

Fisher, W. K. 1928. Asteroidea of the North Pacific and adjacent waters. Pt. 2. Forcipulata.— 
Bulletin of the United States National Museum 76, 245 pp. 

——. 1930. Asteroidea of the North Pacific and adjacent waters. Pt. 3. Forcipulata.—Bulletin of 
the United States National Museum 76, 356 pp. 

Franz, D. R., and A. S. Mermill. 1980. The origins and determinants of molluscan faunal groups on 

the shallow continental shelf of the Northwest Atlantic—Malacologia 19(2):227—248. 

, E. K. Worley, and A. A. Mermill. 1981. Distribution patterns of common seastars of the 

Middle Atlantic Continental Shelf of the Northwest Atlantic (Gulf of Maine to Cape Hat- 

teras).—Biological Bulletin 160:394—418. 

Grainger, E. H. 1966. Seastars (Echinodermata: Asteroidea) of Arctic North America.—Fisheries 
Research Board of Canada, Bulletin 152, 70 pp. 


VOLUME 96, NUMBER 3 547 


Gray, I. E., M. Downey, and M. J. Cerame-Vivas. 1968. Sea stars of North Carolina.—United 
States Fish and Wildlife Service Fisheries, Bulletin 87(1): 127-163. 

Hulbert, A. W. 1980. The functional role of Asterias vulgaris Verrill (1866) in three subtidal com- 
munities.—Ph.D. Thesis, University of New Hampshire, 174 pp. 

Hyman, L.H. 1955. The invertebrates. Vol. 1V. Echinodermata. McGraw-Hill Book Co., Inc., New 
York, 763 pp. 

Kraeuter, J. N. 1974. Offshore currents, larval transport, and establishment in southern populations 
of Littorina littorea along the U.S. Atlantic coast.—Thalassia Jugoslavica 10(1/2): 150-170. 

MacNeil, F. S. 1965. Evolution and distribution of the genus Mya and Tertiary migrations of 
Mollusca.—United States Geological Survey Professional Paper 483-G, 51 pp. 

Mayr, E. 1963. Animal species and evolution.—The Belknap Press of Harvard Univ. Press, 
797 pp. 

McAlice, B. J. 1981. On the post-glacial history of Acartia tonsa (Copepoda: Calanoida) in the Gulf 
of Maine and the Gulf of St. Lawrence.—Marine Biology 64(3):267—272. 

McIntyre, A., et al. [35 others Authors] 1976. Glacial North America 18,000 years ago: A CLIMAP 
reconstruction.—Geological Society of America 145:43-75. 

Mortensen, T. H. 1927. Handbook of the echinoderms of the British Isles —Oxford University 
Press, London, 471 pp. 

Nesis, K. N. 1961. The routes and periods and formation of the interrupted area of distribution of 
amphiboreal species of marine bottom animals.—Okeanologiya 1(5):894—903. 

Schopf, T. J. M., and L. S. Murphy. 1973. Protein polymorphism of the hybridizing seastars 
Asterias forbesi and Asterias vulgaris and implications for their evolution.—Biological Bulletin 
145:587—597. 

Tortonese, E. 1963. Differenziazione geografica e superspecie nel genere Asterias (Echinoderma- 
ta).—Monitore Zoologica Italiano 80—81:212-221. 

Turner, R., and J. H. Dearborn. 1972. Skeletal morphology of the mud star, Ctenodiscus crispatus 
(Echinodermata: Asteroidea).—Journal of Morphology 138(2):239-262. 

Vernill, A. E. 1866. On the polyps and echinoderms of New England with descriptions of new 
species.—Proceedings of the Boston Society of Natural History 10:345-349. 

Vevers, H. G. 1949. The biology of Asterias rubens L. 1. Growth and reproduction.—Journal of 
the Marine Biological Association of the United Kingdom 28:165—187. 

Walker, C. W. 1974. Studies on the reproductive systems of the seastars. I. The morphology and 
histology of the gonads of Asterias vulgaris.—Biological Bulletin 147:661—677. 


Biology Department, Brooklyn College/CUNY, Brooklyn, New York 11210. 


PROC. BIOL. SOC. WASH. 
96(3), 1983, pp. 548-559 


NOTES ON THE FROG GENUS CYCLORAMPAUS 
(AMPHIBIA: LEPTODACTYLIDAE), WITH 
DESCRIPTIONS OF TWO NEW SPECIES 


W. Ronald Heyer 


Abstract.—Examination of the types of Telmatobius duseni Andersson indi- 
cates that a new name is required for the taxon previously referred to as Cy- 
cloramphus duseni. The name Cycloramphus izecksohni is proposed for this tax- 
on. A second new species, Cycloramphus cedrensis, is described for material 
recently collected near Rio dos Cedros, Santa Catarina, Brazil. New distributional 
data are reported for Cycloramphus valae and the advertising call and larva of 
C. valae are described for the first time. 


Information has recently become available that extends our knowledge of the 
systematics and natural history for members of the frog genus Cycloramphus. 
The purpose of this paper is to report these data, which augment those presented 
in a recent revision (Heyer 1983). 


Materials and Methods 


Morphological data were taken and are reported on in a standard manner (e.g., 
Heyer 1983). Advertisement calls were recorded on a Sony TCM-280 cassette 
recorder using a Sennheiser K30 microphone. Calls were analyzed on a Kay 
Sonagraph 6061B, narrow filter, AGC in the off position. Call parameters are 
based on analysis of 5 calls of C. cedrensis and 3 calls each of C. valae from the 
two localities reported. 


The Identity of Telmatobius duseni Andersson 


I had the opportunity to examine two syntypes of Telmatobius duseni after my 
previous study (Heyer 1983) was completed. In that study, the name duseni was 
applied to specimens mostly from the State of Santa Catarina, with one outlier 
locality in the State of Sao Paulo (Heyer 1983: fig. 42). Examination of the types 
of T. duseni indicates they do not represent the species I referred to as duseni. 

The types show that duseni is a member of the C. fuliginosus group; that is, 
the species has toe webbing and dorsal warts and tubercles (Fig. 1). They further 
resemble the species previously referred to as C. duseni and differ from all other 
members of the C. fuliginosus group in small size (types of T. duseni males 31.3, 
33.5 mm SVL), in having a dorsum with shagreen and tubercles with the large 
tubercles sometimes arranged in regular rows (as in Fig. 1), and the posterior 
surface of the thigh with distinct light spots (as in Fig. 1). The types of T. duseni 
differ significantly from the species previously referred to as C. duseni in the 
amount of toe webbing. The types of T. duseni have reduced webbing (webbing 
formula of right foot of lectotype: I trace II 134—3* III 24-34% IV 314-2+ V), the 
species previously referred to as C. duseni has moderate toe webbing. The web- 


VOLUME 96, NUMBER 3 549 


Fig. 1. Lectotype of Telmatobius duseni Andersson, dorsal and ventral views. 


bing formula for the outer side of toe IV in the types of 7. duseni ranges from 
3¥%2 to 4—, whereas webbing in the species previously referred to as C. duseni 
ranges from 24 to 3*, as an example. Such a difference is indicative of species 
level differentiation in the genus Cycloramphus (Heyer 1983). 

At present, the species Cycloramphus duseni (Andersson) is known only from 
the type series. The type locality, ‘“Brazil, Parana, Sierra do mar; Ypiranga, in 
crevices and cracks in the vertical cliffs along the railway, 3/9 1911,’’ (Andersson 
1914:2) was incorrectly plotted in the review paper (Heyer 1983) as discussed in 
Heyer and Maxson (1983). In order to clarify the precise locality where Dusén 
collected the specimens named by Andersson, Dr. P. E. Vanzolini kindly made 
arrangements for me to interview officials of the Rede Ferroviaria Federal S/A 
in Curitiba, Parana. At the railroad office I learned that there is no railway station 
named Ipiranga on the railway line from Curitiba to Paranagua. Rather, Dusén 
collected at Casa Ipiranga, a house on the railway line where the railroad con- 
struction engineers lived during the railroad line construction. The Casa Ipiranga 
today is a state historical preservation site (patrimonio), that can be reached only 
by railroad. Although Casa Ipiranga is not a regular railroad stop, the officials of 
the railway arranged an overnight visit for me on 7 January 1982. Vertical cliffs 
where water was seeping and dripping were found along the railway line. No 
Cycloramphus were found at night along the vertical cliffs or along the stream 
near the house. The habitat looked suitable for Cycloramphus; a longer trip earlier 
in the wet season would likely yield positive results. 

For nomenclatural stability, I hereby designate the 31.3 mm SVL specimen, 
Naturhistoriska Rijksmuseet, Stockholm No. 1606 (Fig. 1), as the lectotype of 
Telmatobius duseni Andersson. The Stockholm Museum has two of the original 
series of five specimens. One specimen was exchanged with the British Museum 
and two specimens are apparently lost (Bengt-Olov Stolt, pers. comm.). Addi- 


550 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


tional characteristics of the lectotype, not apparent in the photograph (Fig. 1) or 
mentioned above are: thumb lacking asperities; snout sloping in profile (Heyer 
1983: fig. 6C); distinct inguinal disk; head length 12.4 mm; head width 13.9 mm; 
eye—nostril distance 2.6 mm; eye—eye distance 5.9 mm; femur length 15.6 mm; 
tibia length 15.8 mm; foot length 16.0 mm. 

Subsequent to my visit to the type locality of duseni at Casa Ipiranga, Werner 
C. A. Bokermann informed me that he had collected at Banhado, the nearest 
regular train station to Casa Ipiranga on the line from Paranagua to Curitiba. I 
had not seen these Cycloramphus previously, so Bokermann loaned them to me 
for purposes of this paper. The specimens (which include two males 37.4 and 
41.4 mm SVL) are C. rhyakonastes, the only known species of Cycloramphus to 
have red bellies in life. Andersson (1914:2) gives the belly color of T. duseni as 
‘‘vellowish dirty white,’ but itis not apparent from the description whether these 
are life colors or those in preservative when Andersson described them. The 
types of T. duseni are similar to Cycloramphus rhyakonastes in dorsal texture, 
posterior thigh surface pattern and foot webbing (the reduced webbing of T. 
duseni is matched by some individuals of C. rhyakonastes). The types of T. duseni 
differ from C. rhyakonastes males in being smaller (males of 7. duseni 31-37 mm 
SVL; males of C. rhyakonastes 37-50 mm). The size difference, probable differ- 
ences in life belly color, and habitat (C. rhyakonastes have only been collected 
along streams, not on rock wall seeps) suggest that duseni and rhyakonastes 
represent two distinct species. Thus I prefer to recognize duseni and rhyako- 
nastes as valid species of Cycloramphus, pending the availability of fresh topo- 
typic material of duseni. 

Cycloramphus duseni (Andersson) will key out to couplet 16 in the key to 
species published previously (Heyer 1983). The smaller size of C. duseni distin- 
guishes it from both C. rhyakonastes and semipalmatus, the species contrasted 
in that couplet. 


A New Name for the Form Previously 
Referred to as Cycloramphus duseni 


The species previously referred to as C. duseni (Heyer 1983; Heyer and Max- 
son 1983) is not the same form represented by the types of Telmatobius duseni 
Andersson. There is no available name for the species, so it is described as: 


Cycloramphus izecksohni, new species 
Fig. 3 


Holotype.—Museu de Zoologia, Universidade de Sao Paulo (MZUSP) 57775, 
an adult male from Brazil; Santa Catarina, 13 km W Pirabeiraba, 26°12’S, 49°07’ W. 
Collected by Annelise Gehrau and W. Ronald Heyer on 17 December 1978. 

Paratopotypes.—MZUSP 57772-74, 57776-98; USNM 217869-96, 217931. 

Referred specimens: All other specimens listed in the distribution section in 
the species account for C. duseni for the States of Santa Catarina and Sao Paulo 
in Heyer (1983). 

Diagnosis.—The species with toe webbing and distinctly black and/or white 
dorsal tubercles that might be confused with the new form are C. asper, bora- 


VOLUME 96, NUMBER 3 551 


46 


22 


28 


52 


Fig. 2. Map of several Cycloramphus distributions in the States of Sao Paulo, Parana, and Santa 
Catarina, Brazil. Base map is adapted from predicted areas of stream associated Cycloramphus 
distributions (Heyer and Maxson 1983). Dashed line is approximate east boundary of the Atlantic 
Forest Morphoclimatic Domain. Stippled areas are zones of sharp relief within the Atlantic Forest 
Morphoclimatic Domain where mountain brooks would be expected. Open circle is previously plotted 
(Heyer 1983) type locality of Telmatobius duseni. Solid circle is correct type locality of T. duseni. 
Triangles are known localities for Cycloramphus izecksoni, new species. Hexagon is only known 
locality for Cycloramphus cedrensis, new species. Squares are known localities for Cycloramphus 
valae. 


ceiensis, cedrensis, duseni, lutzorum, mirandaribeiroi, ohausi, rhyakonastes, and 
semipalmatus. Cycloramphus izecksohni differs from C. ohausi in lacking thumb 
spines. The moderate web of C. izecksohni differentiates it from the forms with 
reduced webbing (C. duseni, rhyakonastes and semipalmatus) or considerable 
webbing (asper, cedrensis and mirandaribeiroi). Cycloramphus izecksohni is 
smaller (males 29-38 mm, females 31-44 mm SVL) than asper, boraceiensis, 
lutzorum, mirandaribeiroi, and rhyakonastes (smallest male 35.9 mm, smallest 
female 42.6 mm SVL). 

Description of holotype.—Snout rounded from above and in profile; canthus 
rostralis indistinct; loreal slightly concave in cross section; tympanum hidden; 


Sp PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 3. Holotype of Cycloramphus izecksohni, dorsal and ventral views. 


vomerine teeth in transverse series almost in medial contact, between and pos- 
terior to choanae; vocal fold present, but vocal slits absent; first finger just shorter 
than second; outer metacarpal tubercle a rounded square with an inner anterior 
projection, inner metacarpal tubercle ovoid, separated from subarticular tubercle 
of thumb by about diameter of subarticular tubercle; thumb lacking asperities; 
dorsal texture with shagreen and small to large warts each bearing a keratinized 
brown or black tubercle; large warts and tubercles arranged in short symmetrical 
rows behind the eyes; weak fold over ear region; inguinal gland large, disk- 
shaped, diameter between 4 and 4% femur length; venter weakly granular; toe 
tips just wider than digits; toes moderately webbed, I 1-27 II 1-2* III 17-24 1V 
242-1 V; toe subarticular tubercles moderately developed; outer metatarsal tu- 
bercle round, smaller than large ovoid inner metatarsal tubercle; no tarsal or 
metatarsal folds; outer tarsus covered with small tubercles, sole of foot smooth. 

SVL 34.9, head length 12.8, head width 14.5, eye—nostril, distance 3.3, eye—eye 
distance 6.6, femur 17.4, tibia 17.3, foot 17.8 mm. 

In preservation, dorsum brown with slightly darker brown amorphous blotches, 
broad lighter interorbital band; loreal and upper lip region with three ill-defined 
light vertical stripes; upper limbs barred; throat, chest, and under sides of limbs 
brown with irregular white spots and marks, belly mostly cream with some brown 
punctations; posterior surface of thigh mostly uniform brown with a few small 
distinct light spots. 

Etymology.—Named for Professor Eugénio Izecksohn, who has contributed 
much to the understanding of the anuran fauna of Brazil and who has consistently 
helped my research efforts. 

The data for adult specimens, advertising call, and larval description are pre- 
sented elsewhere (Heyer 1983, as C. duseni). 

The currently known distribution of C. izecksohni centers on the northern 
portion of the Atlantic Forests in the State of Santa Catarina (Fig. 2). There are 
three specimens of this species (Museu Nacional, Rio de Janeiro 98) with the 
locality data of Alto da Serra, Santos, Sao Paulo. This locality (Fig. 2, locality 


VOLUME 96, NUMBER 3 553 


in State of Sao Paulo) does not make zoogeographic sense according to the scheme 
proposed by Heyer and Maxson (1983). However, there is no reason to doubt 
the veracity of the data and, there is unfortunately little likelihood of verification 
of this species at that locality. In the hundreds of preserved specimens examined 
from the area of Alto da Serra, only one small juvenile was found that might 
represent C. izecksohni (MZUSP 13909). Today, much of the area has been de- 
forested by pollution from steel mills. 


A New Species of Cycloramphus from Santa Catarina 


One of the purposes of the field trip of 1982 was to test the accuracy of pre- 
dictions as to areas of distribution for stream associated Cycloramphus (see Hey- 
er and Maxson 1983), particularly in the southern portion of its range. As ex- 
pected, much of the easily accessible area has been deforested, precluding 
Cycloramphus. We did find a stream with Cycloramphus near the town of Rio 
dos Cedros, Santa Catarina, however. To my knowledge this locality had not 
been sampled previously; examination of the specimens in the laboratory indi- 
cates that they represent a new species, which is named: 


Cycloramphus cedrensis, new species 
Fig. 4 


Holotype.—MZUSP 59260, an adult male from Brazil; Santa Catarina, 12 km 
E of Rio dos Cedros on road to rio Sao Bernardo, approximately 26°44’S, 49°20’W. 
Collected by W. Ronald Heyer on 10 January 1982. 

Paratopotypes.—MZUSP 59256—59, USNM 22978 1-86, same data as holotype, 
except collected on 9 and 10 January 1982. 

Diagnosis.—The other species that share toe webbing and distinct black and/ 
or white dorsal tubercles with the new form are C. asper, boraceiensis, duseni, 
izecksohni, lutzorum, mirandaribeiroi, ohausi, rhyakonastes, and semipalmatus. 
Cycloramphus cedrensis differs from C. ohausi in lacking thumb spines. The 
considerable web of C. cedrensis separates it from C. duseni, rhyakonastes, and 
semipalmatus, which have reduced webbing, and from boraceiensis, izecksohni, 
and /utzorum, which have moderate webbing. Cycloramphus cedrensis is smaller 
(males to 44, females to 50 mm SVL) than mirandaribeiroi (minimum male size 
49, female 62 mm SVL). Cycloramphus cedrensis most closely resembles C. 
asper since both have considerable webbing. Most C. asper are more fully webbed 
than C. cedrensis: modal web formula for female C. asper (female C. asper have 
a little less web than male asper) I 0-27 II 0-2+ III 0-2% IV 2-—0 V, modal web 
formula for cedrensis 1 1-2~ If 1-2% HI 1-2% IV 2%-1 V. The differences in 
webbing, although difficult to verbalize, are apparent when series of specimens 
are compared directly. 

Description of holotype.—Snout rounded from above and in profile, lip flared; 
canthus rostralis indistinct; loreal region slightly concave; tympanum hidden; 
vomerine teeth in almost contiguous transverse series, posterior to choanae; vocal 
slits present; first finger just shorter than second; outer metacarpal tubercle large, 
roughly heart-shaped, inner metacarpal tubercle ovoid, separated from thumb 
subarticular tubercle by distance equal to about 2 diameter of subarticular tu- 
bercle; thumbs lacking asperities; dorsal texture rough, with shagreen and larger 


554 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 4. Holotype of Cycloramphus cedrensis, dorsal and ventral views. 


and smaller warts bearing black or white tipped tubercles, tubercles and warts 
not particularly regularly arranged; fold over ear region distinct; inguinal disk 
large, diameter about 4 length of femur; ventral texture granular; toe tips just 
wider than digits; toes considerably webbed, I 0-2 II 0-2* III 0-2 IV 2!4-0 V; 
rounded outer metatarsal tubercle smaller than ovoid inner metatarsal tubercle; 
no metatarsal fold, a tarsal ridge, but not developed into a fold; outer tarsus with 
many black tipped tubercles; lateral sole of foot with black tipped tubercles, inner 
sole smooth. 

SVL 40.5, head length 16.1, head width 17.3, eye—nostril distance 4.0, eye—eye 
distance 8.3, femur 19.5, tibia 18.7, foot 18.7 mm. 

In preservation, dorsum brown with indistinct tan mottling; upper lip and loreal 
region with three very faint light vertical stripes; upper limbs barred; throat brown 
with white flecks, belly with pattern of small scale brown and white mottle, under 
limbs more or less uniform brown except for light axillary region on arms; pos- 
terior surface of thigh with small regular and irregular light spots. 

Variation.—The 5 adult males and 3 adult females range in size from 40.443 .9 
and 41.9-49.9 mm respectively. The paratypes resemble the holotype in most 
features, but differ in the following features. The posterior surface of the thigh is 
uniform in one specimen, mottled in three others. Light lip stripes are distinct in 
two specimens. The belly is uniform in five specimens. Three individuals have 
the large tuberculate dorsal warts arranged in irregular rows. 

The colors in life of USNM 229784 were: chin and chest with small white dots 
on a maroon background; sides of body and posterior surface of thighs with 
distinctive small yellow dots; iris very dark brown. 

Etymology.—Named for the type locality. 

Advertising call.—Call of sporadic single notes; call short, duration 0.06—0.10 
s; call not well tuned, maximum energy between 1100—1400 Hz, call pulsatile with 
sidebands or harmonics of about 200 Hz (Fig. 5). 


VOLUME 96, NUMBER 3 555 


8 
= 6 
= 
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rT ; 
= f 
m 2 2 
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TIME (S) 


Fig. 5. The advertising call of Cycloramphus cedrensis. Recorded while calling from plastic bag. 
Male individuals in bag were MZUSP 59256-57, 59259-60. Air temperature approximately 27°C. 


Larval definition.—Larvae unknown. 

Distribution.—Presently known only from the type locality (Fig. 2). 

Cycloramphus cedrensis is very similar morphologically to C. asper and the 
two taxa differ only in the degree of webbing. An argument could be made that 
the population described here as C. cedrensis is a geographic variant of C. asper. 
Toe webbing is generally an indicator of species differentiation in the other species 
of Cycloramphus (Heyer 1983; Heyer and Maxson 1983); but the differences in 
webbing are often more pronounced than in this case. The differences between 
C. asper and cedrensis are within the considerably webbed category. The differ- 
ences are consistent and [| interpret them to reflect species differentiation. The 
two species also differ in life color. The bright yellow spots on the sides of the 
body and posterior surfaces of the thighs in C. cedrensis are very distinctive; C. 
asper has no flash colors. Most C. asper are found in the southern half of the 
Atlantic Forest region in the State of Santa Catarina (Heyer 1983, fig. 30). A 
single specimen of C. asper is known from Serra Alta, near Sao Bento do Sul in 
the northern half of the Atlantic Forest region in the State of Santa Catarina 


Table 1.—Size ranges (in mm) of adult Cycloramphus valae from three localities. Numbers in 
parentheses are numbers of individuals. 


Locality Males Females 
Gruta 28.7—32.4 (5) 31.0-39.6 (27) 
nr. Timbé do Sul 30.4—37.3 (10) 34.9-38.4 (3) 


nr. Praia Grande 26. 1-32.1 (7) 29.2—34.0 (7) 


556 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


4 - + 3 
t PP EEG EETEL EE EE EEE EE EE ES. 
N 
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= 
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(S) 
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Fig. 6. The advertising call of Cycloramphus valae. Upper figure from USNM 229814, Brazil; 
Santa Catarina, 15 km from Timbé do Sul on road to Bom Jesus (Rio Grande do Sul), 20:45 h, 
estimated air temperature 18—20° C, 16 January 1982. Lower figure from Brazil; Santa Catarina, 
10 km by road from Itaimbezinho to Praia Grande, 20:10 h, 17—18° C, 14 January 1982, speci- 
men not captured. 


(Adolfo Lutz previously unnumbered specimen 176, Museu Nacional, Rio de 
Janeiro). This specimen has not been reexamined for this study, but the webbing 
formula is consistent with C. asper. 

No other Cycloramphus were found along the stream where C. cedrensis was 
taken. Another species, C. izecksohni has been taken from a nearby locality, 
Timbo do Sul (unnumbered series in E. Izecksohn collection, examined for this 
study) (Fig. 2, triangle next to hexagon). 


New Data for Cycloramphus valae 


Distribution.—The previously documented southernmost record of Cyclo- 
ramphus was for Cycloramphus valae (Fig. 2, northernmost square; the locality 
reported by Braun and Braun, 1980, for Rio Grande do Sul is in error, see Heyer 
1983). The Atlantic Forest Morphoclimatic Domain extends further south than 
this previous locality record; transects across the southernmost extent of the 
Atlantic Forest Domain produced two more records for C. valae (Fig. 2, two 
most southern squares). We did not collect the species in the State of Rio Grande 
do Sul, but further field work is necessary before its absence from the coastal 
mountain range in northern Rio Grande do Sul is accepted. 

There are no noticeable morphological differences among the three available 
geographic samples of C. valae. The southernmost individuals are somewhat 


VOLUME 96, NUMBER 3 IST 


Fig. 7. The larva of Cycloramphus valae, USNM 229809, lateral view and mouthparts. 


smaller in size than the others, although the difference may be due to sampling 
error (Table 1). 

Advertising calls.—The call of C. vaiae is reported here for the first time. Three 
calls each were recorded from two individuals, one from near Timbé do Sul 
(middle square, Fig. 2) and one from near Praia Grande (bottom square, Fig. 2). 
As the calls differ between localities, they are described separately. 

The call of C. valae from near Timbé do Sul consists of 23-26 notes lasting 
from 1.6 to 1.9 s; the dominant frequency ranges between 1250-2250 Hz; each 
note lasts about 0.02 s; the notes are pulsatile, with sidebands or harmonics of 
about 100 Hz; the call intensity is modulated, beginning quietly, ending loudly 
(Fig. 6, top). 

The call of C. valae from near Praia Grande consists of 16-19 notes lasting 
from 1.3 to 1.5 s; the dominant frequency ranges between 1750—2200 Hz; each 
note lasts about 0.02—0.03 s; the notes are pulsatile, with sidebands or harmonics 
of about 80-100 Hz; the call intensity is modulated, beginning quietly, ending 
loudly (Fig. 6, bottom). 

Additional recordings are not available to determine the degree of intraspecific 
variation. The calls of the two individuals available differ in degree of call length 
and number of notes per call. Otherwise, the calls are similar. The similarity of 
calls suggests that the samples are conspecific, although geographic variation may 
prove to be present. 

Larval definition.—Larvae are elongate and depressed (Fig. 7). The tail fins 
are low, present only on the distal half of the tail. The belly is flattened and has 
a shallowly bilobed flap which extends posteriorly past the body. The spiracle, 
if present, is not visible. The anus is median. The eye is moderately large, 16— 
19% of the head—body length. The oral disk is broad, 39-41% of the head—body 
length. The oral disk is entire. The anterior papilla gap is broad, almost the width 
of the disk. The oral papillae are in a single row, continuous posteriorly. The 

1 


tooth row formula is iy the split tooth row halves either abutting or overlapping 


y, 
medially. The beak has strong and deep central cusps. The head—body length is 
24-29% the total length. The largest total length is 29.3 mm of a stage 36 (Gosner 
1960) larva. The body is uniform dark brown above, mottled on the sides, and 


558 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


unpigmented below; the tail ranges from almost uniform brown above to a pattern 
with light dorsal ocelli, the tail venter is unpigmented. 

Larvae were collected from water film covered vertical rock walls of a road 
cut. Of the many tadpoles collected from this habitat at this locality, only three 
were C. valae; the other larvae were all Thoropa. 


Discussion 


The distributional data reported herein do not particularly clarify any zoogeo- 
graphical patterns aside from establishing the southern distributional limit of C. 
valae. In the latter case, Cycloramphus was predicted to occur in the places 
reported above (Heyer and Maxson 1983), and either C. valae or an unknown 
congener would have been expected to occur in the southernmost part of the 
generic range. Resolution of the location of the type locality of C. duseni removes 
one zoogeographic enigma caused by faulty map plotting (Heyer 1983). As pres- 
ently understood, the distributions of C. asper and C. izecksohni are enigmatic 
in that each species has a central range with a geographic outlier population. 
These outlier populations are difficult to explain. 

Cycloramphus valae is a member of the C. granulosus morphological group 
(based on adult morphology, Heyer 1983). The relationships of this group lie with 
the C. fuliginosus group: either the groups are closely related or the granulosus 
group is comprised of ectomorph species, each of which has as its closest relative 
a member of the fuliginosus group (Heyer and Maxson 1983). Neither advertising 
calls nor tadpoles were known for any member of the C. granulosus group pre- 
vious to this report. The larvae share the same derived habitat and morphology 
as the known larvae of the C. fuliginosus group; there is nothing in the larval 
features to suggest that C. valae differs significantly from members of the fuligi- 
nosus group. The advertising call of C. valae differs in kind from the calls known 
for other members of the fuliginosus group. The calls of the fuliginosus group 
consist either of three note calls, the first note differing from the final two, or 
single note calls (which are similar to the final two notes of the three note call) 
(Heyer 1983). The call of C. valae, in contrast, consists of a multi-note call, 
similar in overall call pattern to that of C. ohausi, the sole representative of the 
C. ohausi group. The individual notes of the C. valae call are similar to notes of 
fuliginosus group members in having harmonics or sidebands. The individual 
notes of C. ohausi lack these features. The call of C. valae thus combines com- 
ponents found in calls of members of other species groups. Conclusions regarding 
relationships based on calls must await analysis of this feature in other members 
of the granulosus morphological group. 


Acknowledgments 


Bengt-Olov Stolt (Naturhistoriska Rijksmuseet, Stockholm) facilitated the loan 
of the types of Telmatobius duseni. Srs. Denisar Zanello Miranda and Joao Fer- 
nando Corsico, Rede Ferroviaria Federal S/A, Curitiba, most graciously arranged 
the trip to Casa Ipiranga. Francisca Carolina do Val (Museu de Zoologia, Uni- 
versidade de Sao Paulo, MZUSP) made the field work possible and enjoyable. 
P. E. Vanzolini (MZUSP) clarified points concerning localities and collectors and 
reviewed the manuscript. Frances Irish prepared Fig. 7. Ronald I. Crombie and 


VOLUME 96, NUMBER 3 559 


George R. Zug (National Museum of Natural History, USNM) also reviewed the 
manuscript. Field work was supported by the International Environmental Sci- 
ences Program, Smithsonian Institution, and the Museu de Zoologia da Univer- 
sidade de Sao Paulo. 


Literature Cited 


Andersson, L. G. 1914. A new Telmatobius and new Teiidoid lizards from South America.—Arkiv 
for Zoologi 9: 1-12. 

Braun, P. C., and C. A. S. Braun. 1980. Lista prévia dos anfibios do Estado do Rio Grande do Sul, 
Brasil.—Iheringia 56:121-146. 

Gosner, K. L. 1960. A simplified table for staging anuran embryos and larvae with notes on iden- 
tification.—Herpetologica 16:183—190. 

Heyer, W. R. 1983. Variation and systematics of frogs of the genus Cycloramphus (Amphibia, 
Leptodactylidae).—Arquivos de Zoologia 30:235-339. 

Heyer, W. R., and L. R. Maxson. 1983. Relationships, zoogeography, and speciation mechanisms 
of frogs of the genus Cycloramphus (Amphibia, Leptodactylidae).—Arquivos de Zoologia 30: 
341-368. 


W. Ronald Heyer, Amphibians and Reptiles, Smithsonian Institution, Wash- 
ington, D.C. 20560. 


PROC. BIOL. SOC. WASH. 
96(3), 1983, pp. 560-566 


LEPTODACTYLUS RIVEROI, A NEW FROG SPECIES 
FROM AMAZONIA, SOUTH AMERICA 
(ANURA: LEPTODACTYLIDAE) 


W. Ronald Heyer and William F. Pyburn 


Abstract.—A new species of the frog genus Leptodactylus is described based 
on specimens from scattered localities in Amazonia. The new species has char- 
acteristics which bridge the morphological gap between the previous definitions 
of the melanonotus and ocellatus species groups. 


Rivero (1968) recognized that a collection of 27 individuals of Leptodactylus 
from Cano Iguapo, Venezuela represented a distinctive species of Leptodactylus, 
different from those he had treated previously in the “‘Salientia of Venezuela’ 
(1961). In the 1961 publication, Rivero questionably included the species L. rho- 
domystax in the Venezuelan fauna, based on a single juvenile in the collections 
of the American Museum of Natural History. At the same time he pointed out 
that Boulenger’s (1883) description of L. rhodomystax was based on a juvenile. 
Rivero later (1968) concluded that the series of specimens from Cano Iguapo 
represented L. rhodomystax Boulenger. Boulenger’s L. rhodomystax is a differ- 
ent species, however (Heyer 1979), leaving the species Rivero recognized as 
distinct without a name. In recognition of Dr. Rivero’s contributions, we describe 
the species as 


Leptodactylus riveroi new species 
Fig. | 


Holotype. —USNM 232400, an adult male from Colombia; Vaupes; Timbo, 
01°06'S, 70°01'W, elevation 170 m. Collected by William F. Pyburn and J. K. 
Salser, Jr., 25 May 1973. 

Paratypes.—UTACV 3888-3898 (Colombia; Vaupes; Timbo); UTACV 3721 
(Colombia; Vaupes; Wacara); UTACV 3792, 4295, 4319, 6025, 7971-7974 (Co- 
lombia; Vaupes; Yapima). 

Referred specimens.—All non-Colombian specimens listed in the distribution 
section. 

Diagnosis.—Leptodactylus riveroi has extensive toe fringing and thumb spines 
in the males, characteristics shared with members of the Leptodactylus melano- 
notus and ocellatus groups. Leptodactylus riveroi has a pair of low dorsolateral 
folds, indicated in preservative by a black border, extending from behind the eye 
to a point near the posterior end of the body above the groin; L. dantasi, melano- 
notus, podicipinus, pustulatus, and most wagneri lack dorsolateral folds. Some 
L. wagneri have short indications of dorsolateral folds, but they are never as 
extensive as those of L. riveroi. The light stripe from under the eye to the shoul- 
der, uniformly found in L. riveroi, is not present in L. dantasi, melanonotus, 
ocellatus, podicipinus, pustulatus, and most wagneri. Some L. wagneri have a 
light stripe under the eye to the angle of the jaw, but the outlines of the stripe 


561 


VOLUME 96, NUMBER 3 


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562 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


are usually vague and the stripe is not as clearly defined as in L. riveroi. Lep- 
todactylus bolivianus often have distinct light canthal stripes, but the stripes begin 
well forward of the eye, not under it as in L. riveroi. The uniform or faintly 
blotched dorsal pattern and single pair of dorsolateral folds of L. riveroi differ 
from the distinctly spotted dorsal patterns and at least 4 dorsolateral folds of L. 
chaquensis, macrosternum, ocellatus, and viridis. Male L. riveroi further differ 
from all other Leptodactylus (males of L. dantasi not known) in lacking vocal 
slits. 

Specimens of L. riveroi are most likely to be confused with L. bolivianus, 
rhodomystax, and wagneri. The light lip stripes of L. riveroi and L. rhodomystax 
are similar, but L. rhodomystax has no toe fringing in adults and the back of the 
thigh has distinct, discrete light spots, rather than the mottling found in L. riveroi. 
In addition to the differentiating characteristics listed above for bolivianus and 
wagneri, L. bolivianus lacks the red-orange ventral glands found in L. riveroi and 
most adult L. wagneri are smaller than adult L. riveroi. 

Description of holotype.—Snout subovoid from above, rounded in profile; can- 
thus rostralis rounded; lores concave in cross section; tympanum large, distinct, 
almost same diameter as eye diameter; no vocal slits; vomerine teeth in two 
arched series, approaching each other medially, posterior and medial to choanae; 
first finger just longer than second, second about equal to fourth, third much 
longer than others; fingers with lateral fleshy ridges; thumb with 2 cornified nuptial 
spines; arm not especially hypertrophied; no ulnar ridge; dorsal texture smooth; 
a pair of low dorsolateral folds, demarcated by black pigment laterally, folds 
extending from back of eye to back of body above groin, moderate supratympanic 
fold present; ill-defined parotoid glands present, sides of body generally glandular, 
red-orange glands in large blocks on throat, most of belly, and ventral surfaces 
of limbs; venter smooth, ventral disk fold indistinct; no chest spines; toe tips not 
expanded; toe fringe extensively developed; subarticular tubercles moderately 
developed; extensive metatarsal flap of skin; tarsal fold distinct, extending about 
% length of tarsus, just not continuous with toe fringe of first toe; upper tibia 
with many white tipped tubercles; posterior surface of tarsus and sole of foot 
with many black tipped tubercles. 

Snout—vent length (SVL) 62.8 mm, head length 25.0 mm, head width 23.2 mm, 
interorbital distance 5.5 mm, eye—nostril distance 7.1 mm, femur 26.7 mm, tibia 
29.9 mm, foot 35.8 mm. 

Dorsum essentially uniformly brown (in preservative) between dorsolateral folds 
and in upper snout area; faint darker interorbital mark, straight anteriorly, indis- 
tinct posteriorly; two indistinct dark central blotches; dark, broken canthal stripe 
from nostril to eye; upper lip indistinctly marked, lighter area under eye becoming 
distinct light strip under tympanum, extending to beyond angle of jaw, ending in 
shoulder region; sides of body brown with darker spots above, with a lighter 
indistinct band between side and belly; upper limbs indistinctly barred; ventral 
surfaces boldly mottled with dark and light, overlain by red-orange glandular 
areas, posterior surface of thigh boldly mottled with red-orange and black-brown. 

Variation.—Rivero (1968) gave the range of sizes of his sample as 29.0—72.5 
mm, but did not indicate ranges of males and females within that sample. In the 
specimens we have examined, males range 52.2—62.8 mm and females 67.8-81.0 
mm SVL. The ranges of body proportions among adults are: head length/SVL 


VOLUME 96, NUMBER 3 563 


a) 


AANA ha A 
AA NAAN Nitin RAN A , AY: 

ws) i} [VV VV VAVAA aaa AA Aca AUT Wen Aen ene 
Vy Wye ye 


V 


SE ee ee EOE ELEC RCE Rae TEED ELT MeO En 
<2 : Ge Sa et ee 
(MMM MEHU TN yt Loe) np nQMPO NANT) eb poy WaT TT PUT Te TTT Gt Wali Ty 
ie eens momi neous dua ean 


a —$—$$$-_- sss == 


voi Ww i W WW Ww mM We vv We wd wy Ww We we Ww vw we oo av Ww Ww “ Ww Ww we ww 
“ij i 
l 2 
TIME IN SECONDS 


Fig. 2. Advertisement call of Leptodactylus riveroi. Wave form of filtered call as shown in top 
audiospectrogram, showing end of one note and two complete notes. Length of entire signal is 0.214 
s. Upper audiospectrogram of bandpass filtered call, % octave at 500 Hz. Note that time axis is twice 
that of lower audiospectrogram. Lower audiospectrogram unfiltered call. All recordings from Colom- 
bia: Vaupes; Timbo, recorded on 28 May 1973 at about 21:30 h by W. F. Pyburn and J. K. Salser, 
Jr. Water temperature 24.0°C, air temperature 25.8°C. Specimen UTACV 3890 calling from burrow 
in a Swamp. 


38-48%, head width/SVL 35-37%, interorbital distance/SVL 8—10%, eye—nostril 
distance/SVL 10-11%, femur/SVL 40-46%, tibia/SVL 46-48%, and foot/SVL 53- 
57%. 

There is little intraspecific variation in pattern or morphology among the indi- 
viduals at hand. In some of the specimens, the tarsal fold is continuous with the 
outer toe fringe of toe one. 

Rivero (1968) gave the following color notes from the Venezuelan individuals: 
dorsolateral folds at times maroon or cream red; the line that goes from the eye 
to the shoulder, and also at times the loreal area (although never as pronounced) 
ranges from a shade of rose to at times reddish. Charles W. Myers provided the 
following color data (also on Venezuelan individuals) through his notes and a 
color transparency: dorsal coloration basically brown, dorsolateral fold black 
outlined; the stripe under the eye cream anteriorly, orangish posteriorly in shoul- 
der region; rear of thigh mottled black and yellowish brown; ventral surfaces 
mottled yellowish white and light brown; iris pale bronze, with reddish brown 
horizontal stripe, lower part of iris below stripe darkened by heavy black vena- 
tion. 

In life, specimens from southeastern Colombia are dark purplish brown on the 
dorsum and sides. Black bands cross the upper surface of the forearms and thighs, 
and irregular black spots occur on the upper surfaces of the shanks and on the 
sides of the body. Some specimens have a row of black spots above the groin 
that is partially concealed by the leg when the frog is at rest. Black reticulations 
or mottlings cover the posterior thigh surfaces and there is an interocular black 


564 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 3. Known distribution of Leptodactylus riveroi. Political boundaries indicated for Brazil, 
Colombia, and Venezuela. 


bar. A bright cream lip stripe begins below the eye and extends posteriorly along 
the lower edge of the tympanum to the base of the upper arm. As the lip stripe 
approaches the arm base, it may gradually turn to orange. The lore is creamy 
gray, somewhat darker than the lip stripe. The edge of the upper lip is black 
crossed by one or two narrow cream bands. The dorsolateral fold is gray-bronze 
and its lower edge is marked by a broken or continuous black line. Large orange- 
yellow glandular areas cover most of the ventral surfaces, which are also marked 
with dark gray reticulations. 

Advertisement call.—The call is quiet and consists of approximately 9-28 notes 
with a duration of from 0.7—2.3 s. Each note consists of two major pulses and 
the entire note has a duration of 0.04—0.05 s. The dominant frequencies range 
from 360-750 Hz to 360-830 Hz. There appears to be frequency modulation within 
each major pulse and the note is intensity modulated with the second pulse the 
loudest. There is no harmonic structure (Fig. 2). 

Distribution.—The species is thus far known from the following localities 
(Fig. 3): 


VOLUME 96, NUMBER 3 565 


Brazil: Amazonas; Manaus (Reserva Ducke), MZUSP 50170, Reserva INPA- 
WWE (Rio Preto), MZUSP 57966. 

Colombia: Vaupes; Timbo, USNM 232400, UTACV 3888-3898, Umuna (un- 
catalogued voucher specimen), Wacara, UTACV 3721, Yapima, UTACV 3792, 
4295, 4319, 6025, 7971-7974. 

Venezuela: Amazonas; Upper Rio Orinoco, Cano Iguapo (Rivero reported 
specimens, not examined by us), Upper Rio Orinoco, Cano Cotua (between Rio 
Orinoco and Cerro Yapacana), 100 m, AMNH 100655, Upper Rio Orinoco, SW 
base Cerro Yapacana, 110 m, AMNH 100654, E of Purunama on Rio Guaname, 
USNM 229779-780. 

Habitat.—AMNH 100654 was taken at night on the bank of a rocky stream in 
humid evergreen forest (not subject to flooding); AMNH 100655 was collected at 
night on the bank of Cano Cotua in an area of seasonally inundated low scrubby 
forest (caatinga amazonica as used by Venezuelan botanists). At this latter hab- 
itat, L. wagneri was sympatric with L. riveroi. 

In the forest of southeastern Colombia, Leptodactylus riveroi occurs at night 
along the edges of streams and on high ground in swamps that have been formed 
by flooding. When disturbed, it leaps directly into the water, but may turn and 
swim back to the bank where it emerges onto the land and sits quietly among 
dead leaves. In daylight, L. riveroi is occasionally found along forest trails away 
from water. 

J. K. Salser, Jr. and the junior author found a calling male (USNM 232400) 
about 2100 h, 25 May 1973 by locating the source of the sound the animal pro- 
duced. The frog called from the concealment of an underground cavity connected 
to the surface by a slanting tunnel near the edge of a swamp. The cavity was 
intersected by roots and contained a small pool. A regular sequence of quiet, 
closely spaced, low-pitched notes emanated from the cavity. 

Another male (UTACV 3890) recorded (Fig. 2) 28 May 1973 at the same locality 
called from an underground chamber essentially like that of USNM 232400. The 
chamber was covered over by roots, dead leaves and humus. It contained a pool 
of water 25 cm deep and was located in a hillock surrounded by the water of the 
swamp. An adult female (UTACV 3891) sat on another hillock 2 m from the 
calling male. 


Discussion 


Leptodactylus riveroi demonstrates characters that straddle the previously de- 
fined L. melanonotus and ocellatus groups. The only characteristic that separated 
the two groups was the presence of well defined dorsolateral folds in the ocellatus 
group. Thus, on this basis, L. riveroi would be a member of the ocellatus group. 
However, the species has an overall habitus and color pattern strikingly similar 
to L. wagneri, a member of the melanonotus group. The unique condition in L. 
riveroi of having no vocal slits in males confuses the question of the precise 
relationships of L. riveroi. The call structure of L. riveroi stands out in its dis- 
tinctiveness and is unlike any other Leptodactylus known. At the least, L. riveroi 
shows that there is no clear cut morphological distinction between the L. melano- 
notus and ocellatus groups. An albumin sample of L. riveroi is available and the 


566 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


micro-complement fixation analyses of Leptodactylus albumin including the sam- 
ple of L. riveroi will be reported elsewhere (Maxson and Heyer, in prep.). 


Acknowledgments and Museum Specimen Citation Symbols 


We wish to thank Charles W. Myers, the American Museum of Natural History 
(AMNH) and P. E. Vanzolini, Museu de Zoologia da Universidade de Sao Paulo 
(MZUSP) for the loan of specimens in their charge. Dr. Myers also kindly made 
available his color and habitat notes of L. riveroi. We are indebted to J. K. Salser, 
Jr. and Nathan Waltz for their help in collecting and recording the specimens 
from Timbo and Yapima. Other specimens are deposited in the National Museum 
of Natural History (USNM) and the University of Texas at Arlington Collection 
of Vertebrates (UTACV). Field work by the junior author was made possible by 
a grant from the University of Texas at Arlington Organized Research Fund. 
Funds for publication of the color plate were provided by a Smithsonian Fluid 
Research Grant. 


Literature Cited 


Boulenger, G. A. 1883. Ona collection of frogs from Yurimaguas, Huallaga River, northern Peru.— 
Proceedings of the Zoological Society of London 1883:635-638. 

Heyer, W.R. 1979. Systematics of the pentuductylus species group of the frog genus Leptodactylus 
(Amphibia: Leptodactylidae).—Smithsonian Contributions to Zoology 301: 1-43. 

Rivero, J. A. 1961. Salientia of Venezuela.—Bulletin of the Museum of Comparative Zoology 126: 
1-207. 

——. 1968. El problema de Leptoductylus rhodomystux Boulenger (Amphibia, Salientia)—Me- 
moria de la Sociedad de Ciencias Naturales La Salle 28:145—-150. 


(WRH) Amphibians and Reptiles, National Museum of Natural History, Smith- 
sonian Institution, Washington, D.C. 20560; (WFP) Department of Biology, Uni- 
versity of Texas at Arlington, Arlington, Texas 76019. 


PROC. BIOL. SOC. WASH. 
96(3), 1983, pp. 567-580 


AN ANNOTATED CHECKLIST OF THE FOSSIL TORTOISES 
OF CHINA AND MONGOLIA 


Charles R. Crumly 


Abstract.—A review of published descriptions in some unavailable or obscure 
journals was done to collate and summarize useful morphological data on the 
fossil tortoises of Mongolia and China. Four genera of testudinids are recognized 
and tentatively diagnosed: Testudo, Geochelone, Indotestudo, and ‘Manouria.’ 
Sinohadrianus and Kansuchelys are considered synonyms of ‘Manouria,’ which 
is diagnosable, but not defined by shared derived characters. 


Fossil land tortoises (Testudinidae) appear suddenly in Early Eocene sediments 
of North America and China. By the Late Eocene tortoises are also known from 
Europe (de Broin 1977; Mlynarski 1955; Schleich 1981) and North Africa (An- 
drews 1906), while persisting in North America and China. Fossil tortoises are 
among the most commonly encountered Tertiary remains and may be employed 
as useful stratigraphic markers (Hutchison 1980). 

Although much is known of the structure, distribution, and evolutionary rela- 
tionships of European and North American tortoises (Auffenberg 1963, 1966, 
1971, 1974, 1976; Bramble 1971, in press; Van Devender et al. 1976), compara- 
tively little is known of their Chinese counterparts. The acquisition of detailed 
information on the 24 named Chinese taxa has been hampered by two factors. 
Firstly, is the difficulty in examining the material. During the past forty or so 
years there has been very little interchange between oriental and occidental pa- 
leobiologists. Travel to the Institute of Vertebrate Paleontology and Paleoanthro- 
pology in China, where 17 of the 24 types reside, has not been possible until 
recently. Thus, western students of tortoise evolution, familiar with North Amer- 
ican and European fossil remains have not had the opportunity to examine Chinese 
material. Secondly, the original descriptions of Chinese fossil land tortoises have 
often been published in journals not readily available to western researchers. The 
purpose of this checklist is to summarize the information published in those 
journals, to facilitate a useful synthesis. 

The following list is arranged alphabetically by species name. Comments on 
materials, other than American Museum of Natural History specimens, are based 
on published figures and diagnoses. For this reason and because some of the 
figures are poor, remarks concerning morphology are often incomplete. The pres- 
ent status of the species and prevailing theories of relationship, if any, are noted. 
Generic names in quotes indicate that I have been unable to corroborate mono- 
phyly of the genus. The obvious limitations inherent in a checklist like this require 
that the reader regard generic assignments as tentative. Specimens in the Amer- 
ican Museum of Natural History, New York (AMNH) were examined. Other 
unexamined material is housed at the Institute of Vertebrate Paleontology and 
Paleoanthropology, Beijing (IVPP), Zaklad Paleobiologii, Polska Akadamia Nauk, 
Warszawa (Z. Pal.), and the Geological Survey of China. The bone and scute 


568 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


terminology of Zangerl (1969) and the Catalogue of American Amphibians and 
Reptiles is followed. 


Kansuchelys chiayukuanensis Yeh, 1963a:28 


Type.—IVPP V.1006, right posterior carapace and plastron, damaged. Two 
paratypes are designated (IVPP 1006a, IVPP 1006b). Type locality: Shih-erh-ma- 
ch’eng, North of Hui-hui-p’u, Chia-yu-kuan, Kansu. Horizon: Unknown (? Oli- 
gocene or Late Eocene). 

Remarks.—A very primitive tortoise, judging from the entirely hexagonal neur- 
als, which are short-sided anteriorly, and the primitive suprapygal pattern (two 
suprapygal elements with the larger dorsal element embracing the ventral elliptical 
element. Extended discussion of this and other features will be the subject of 
another paper). Numerous other features show resemblances to ‘Manouria’ (sen- 
su Auffenberg 1971) species. These include the anal scute which is about the 
same median length as the femoral scute and the pleural scutes which are con- 
tacted by only the fifth and sixth marginals. The gular scutes but not the pectoral 
scutes overlap the entoplastron. The cervical scute is very large but slightly longer 
than broad. 


Testudo chienfutungensis Yeh, 1963a:44 


Type.—IVPP V.1030, carapace with connected plastron, damage to anterior 
carapace, posterior peripherals missing. Type locality: Taben-buluk, Tunhuang, 
Kansu. Horizon: Miocene (?). 

Remarks.—Y eh (1963a) allocated this species to the genus Testudo on the basis 
of the position of the pleuro-marginal sulcus. Both Yeh (1963a) and Bohlin (1953) 
felt that the elongate shell, the presence of intergular scutes and the hexagonal 
neurals were abnormal conditions. Irrespective of the presence or absence of 
intergular scutes, the gular scutes overlap the entoplastron. Auffenberg (1974) 
thought this species may not be a testudinid. Bohlin (1953) noted two suprapygals 
arranged serially and of about equal size, whereas Yeh (1963a) characterized the 
suprapygal as entire. The four-sided fourth neural is odd for a tortoise, in which 
this neural is usually eight or six-sided. This species does possess some tortoise- 
like features, but most are shared primitive features. For example, the anal scute 
is much broader than the femoral scute and there is very little constriction at the 
anal and gular scutes posterolateral borders. Auffenberg’s (1974) reservations 
concerning this species are understandable. 


Testudo demissa Gilmore, 1931:239 


Type.—AMNH 6670, right xiphiplastron (probably of large male). Type local- 
ity: Ardyn Obo, Chinese Postroad, Outer Mongolia. Horizon: Ardyn Obo for- 
mation, Lower Oligocene. 

Remarks.—This species is distinguished on the basis of a thickened and down- 
turned anal region of the plastron and very short anal scutes. Both these char- 
acters appear in many unrelated tortoises and are even more prominent in large 
males of these unrelated species. Mlynarski (1968) synonymized 7. demissa with 
Geochelone insolitus (Matthew and Granger, 1923), and allocated both species 


VOLUME 96, NUMBER 3 569 


to the genus Geochelone. Auffenberg (1974) tentatively allocated G. insolitus and 
T. demissa to the subgenus Manouria. The extreme narrowness of the anal scutes 
contradicts Auffenberg’s allocation. 


Testudo hipparionum Wiman, 1930:41 


Type.—IVPP (Catalogue number not published), complete shell. Type locality: 
Shansi, Wu-Hsiang-Hsien, E 2 li (1 li equals about one-third of a mile) Huo- 
Shen-Miao-Kou. Horizon: Pliocene. 

Remarks.—Very poorly described by Wiman (1930) but re-described and fig- 
ured from non-type material by Yeh (1963a). Yeh (1963a) also refers four IVPP 
specimens to this species (IVPP V.1017, V.1018, V.1020, and V.1024). IVPP 
V .1024 includes a skull and parts of the appendicular skeleton. Yeh (1963a) noted 
that this may be the most common tortoise in northern Chinese red clay beds. 
Gilmore (1934) and Auffenberg (1974) contended that other tortoises named by 
Wiman may be variants of 7. hipparionum. The shape and size of the cervical 
and questionably interpreted pleuro-marginal scute contacts support the alloca- 
tion of this species to the genus Testudo (sensu stricto). Other characters include 
two suprapygal elements, the larger embracing the smaller elliptical element 
(primitive); the gular scutes, but not the pectoral scutes, overlap the entoplastron; 
some plastral constriction at the anal and epiplastral border; three marginals 
(numbers 5, 6, and 7) contact the second pleural scute (primitive); and neural 
formula 4-8-4-8-4-8-?4-26. 


Testudo honanensis Wiman, 1930:43 


Type.—IVPP (Catalogue number not published), partial carapace and nearly 
complete plastron. Type locality: locality 12, Honan, Hsin-An-Hsien, Shan-Yin 
Kou. Horizon: Unknown (? Miocene). 

Remarks.—Y eh (1963a) referred three IVPP specimens (IVPP V.1025, V.1026, 
V.1027) to this species but Auffenberg (1974) contended that Yeh’s usage of this 
name is synonymous with 7. sphaerica. Mlynarski (1955) believed that this form 
is assignable to the antiqua-graeca phyletic line. Cervical shape and size, and an 
anal scute having a greater median length than the femoral scute support the 
allocation of this species to the genus 7estudo. The hexagonal third neural is 
unusual. The neural formula is 4-8-6-8-?4-6-6-6. 


Testudo hypercostata Wiman, 1930:35 


Type.—IVPP (Catalogue number not published), nearly complete shell. Type 
locality: locality 114 south, Shansi, Ho-Ch’u-Hsien, SE 60 li (1 li equals about 
one-third of a mile) of Nan-Skha-Wa, | li from Ma-Hua-T’an. Horizon: Pliocene. 

Remarks.—Yeh (1963a) contended that this species is specialized because it 
possesses a supernumerary neural. Auffenberg (1974) disagreed and claimed that 
this form is conspecific with T. shensiensis. The neural formula 1s 4-8-4-8-4-6-6- 
6-4. The anal midline length is only slightly less than the femoral length. The 
entoplastron is not overlapped by either the gular or pectoral scutes. There is 
some constriction at the lateral borders of the epiplastral and anal regions. The 
second pleural scute is contacted by three marginal scutes. 


570 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Testudo insolita Matthew and Granger, 1923:5 


Type.—AMNH 6275, parts of carapaces and plastra of several individuals. 
Type locality: Promontory Bluff, on the Sair-Usu-Kalgan trail, about 150 miles 
from Sair-Usu and 350 miles from Kalgan, Ardyn Obo basin, Mongolia. Horizon: 
Oligocene. 

Remarks.—Gilmore (1931) felt the poor description of this species made the 
determination of its affinities impossible. Mlynarski (1968), nonetheless, synon- 
ymized Geochelone demissa (Gilmore, 1931) with 7. insolitus. The entoplastron 
is not overlapped by the pectoral scutes, which narrow abruptly as they extend 
medially. The xiphi-hypoplastral sutures are located far posterior from the fem- 
oral-abdominal sulcus. The cervical scute is long and thin and recessed from the 
anterior margin of the carapace. Mlynarski’s (1968) assertion that the neural bones 
are octagonal, hexagonal and tetragonal cannot be confirmed until fossil material 
of that region of the shell is found. This large tortoise with abbreviated anal scutes 
is referable to Geochelone, as Auffenberg (1974) and Mlynarski (1968) suggest. 

Chkikvadze (1972) described Ergilemys, type-species G. insolitus, and consid- 
ered that Early Oligocene genus ancestral to Protestudo Chkikvadze (1970), 
‘Manouria,’ Geochelone, and Indotestudo. The distinctive features of Ergilemys 
are primitive characteristics. Mlynarski (1976) demotes Ergilemys to a subgenus 
of Geochelone. In addition to the five species assigned to Ergilemys by Chkik- 
vadze, de Broin (1977) has included another newly named species. Thus, although 
its affinities are often considered, 7. insolita remains enigmatic. 


Testudo kalganensis Gilmore, 1931:247 


Type.—AMNH 6701, anterior part of carapace, most of anterior lobe of plas- 
tron minus entoplastron, and carapacial and plastral parts of right side of bridge. 
Type locality: Kalgan Area, North China. Horizon: Tertiary (?). 

Remarks.—Gilmore (1931) did not remark upon the relationships of this species; 
such reticence is understandable given the greatly thickened and extraordinarily 
sculptured shell. Auffenberg (1974), citing two of his earlier works (1962 and 
1963), assigned this species to the turgida phyletic line of the subgenus Hesper- 
otestudo. The pectoral scute overlaps the entoplastron, a feature that character- 
izes Indotestudo and Testudo horsfieldii. The gular scutes also overlap the en- 
toplastron. The cervical is long and thin and slightly recessed. Although 
questionable, it is possible that the femoral scutes do not contact the inguinal 
scutes, a feature common to Testudo. Assignment to Hesperotestudo seems pre- 
mature. 


Testudo kaiseni Gilmore, 1931:236 


Type.—AMNH 6352, nearly complete carapace and plastron. Type locality: 
Ardyn Obo basin, Chinese Postroad, Outer Mongolia. Horizon: Ardyn Obo for- 
mation, Lower Oligocene. 

Remarks.—Auffenberg (1974) questionably referred this species to Indotestu- 
do, probably because the pectoral scute overlapped the entoplastron. Earlier, 
Glaessner (1935) contended that 7. kaiseni was a member of the Testudo antiqua 


VOLUME 96, NUMBER 3 571 


group, contrary to Mlynarski (1955). The femoro-abdominal sulcus, which is faint- 
ly visible on the type, was not figured by Gilmore (1931, fig. 17, p. 238). The anal 
scute is about the same median length as the femoral scute, perhaps slightly 
greater. Neither the gulars nor the pectorals overlap the entoplastron. The neural 
formula is 4-7-4-7-5-6-?6-?. In most tortoises, the neural series includes neurals 
with an even number of sides. Thus, the seven-sided neurals would probably be 
eight-sided and the five-sided neural would probably be four-sided in other spec- 
imens of Testudo kaiseni. 


Testudo lunanensis Yeh, 1963a:50 


Type.—IVPP V.1032, damaged plastron with left plastral bridge. Type locality: 
Wa-ya-chung, Ta-i-ma, Lunan, Yunnan. Horizon: Early Oligocene. 

Remarks.—Yeh (1963a) distinguished this species, which is the largest tortoise 
known from China, primarily on the basis of size. He believed it to be closely 
related to 7. yuwnnanensis, the other giant tortoise preserved in Chinese deposits. 
Auffenberg (1974) referred this species to the synonymy of 7. yunnanensis. With 
reservation Auffenberg allocated this latter species to Geochelone, on the basis 
of its large size. Only the inside of the plastron, which possesses no particularly 
diagnostic features has been prepared; it is a very poorly known form in which 
size provides the only clue to its affinities. Nevertheless, Chkikvadze (1972) al- 
located this species to his new genus Ergilemys. This species is also represented 
by appendicular material. 


Testudo nanus Gilmore, 1931:241 


Type.—AMNH 6692, complete shell. Type locality: East Mesa, Shara Marun 
Region, Inner Mongolia. Horizon: ? Ulan Gochu, Lower Oligocene. 

Remarks.—Gilmore (1931) anticipated the nomenclatorial change adopted by 
Loveridge and Williams (1957) in noting the mobile posterior plastral lobe in 
Testudo graeca, T. nanus and some other Testudo. In T. nanus, the supracaudal 
scute is divided, but whether it is divided on both the inside and outside of the 
pygal bone is not known. In Geochelone emys and G. impressa it is divided on 
both sides, but in 7. hermanni and Malacochersus tornieri it is usually divided 
only on the outside of the shell. Other characteristics of 7. nanus include an 
elongated prominent epiplastral projection, gular scutes that do not overlap the 
entoplastron which is in turn broadly overlapped by the pectoral scutes, two 
suprapygals with the smaller elliptical ventral element embraced by the larger 
superior element, and no constriction at the anal border. Gilmore (1931) was not 
able to locate the anal/femoral sulcus. However, detailed examination of the type 
material and examination of photos and Gilmore’s plates suggests that the anal 
scute is broader than the femoral; the typical condition for Testudo. Only the pos- 
terior neural bones are known: ?-?-?-?-4-6-6-6. 

Mlynarski (1968) was uncertain about the status of this species but was con- 
vinced that it is not referable to Indotestudo. Auffenberg (1974) allocated this 
species to Geochelone, subgenus questionably Indotestudo. I return this species 
to Testudo because of the mobile plastral lobe and the apparent large size of 
the anal scute. 


Si PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Geochelone oskarkuhni Mlynarski, 1968:91 


Type.—Z. Pal. MgCh/15, a plastron and a pygal region from the same individ- 
ual. Type locality: Altan, Teli, Dzereg Valley, Western Mongolia. Horizon: Plio- 
cene (fide Gradzinski et al. 1968:75). 

Remarks.—The allocation of this species to Geochelone was accepted with 
reservation by Auffenberg (1974). Mlynarski (1968) noted similarities between G. 
oskarkuhni, and Testudo sphaerica Wiman and T. kegenica Khosatsky (1953). 
He also noted that material of G. ulanensis was in too poor a condition to allow 
meaningful comparisons. In order to express his ideas of phyletic relationships 
for Asian tortoises, Mlynarski coined the term “‘hipparionum’’ group and placed 
G. oskarkuhni within this group. He noted that this species group may be inter- 
mediate between Geochelone and Testudo and perhaps ancestral to the antiqua- 
graeca phyletic line. 

The gulars, but not the pectoral scutes, overlap the entoplastron. The cervical 
scute is long and narrow and there are two suprapygal elements. The larger 
superior element embraces the more ventral, elliptical element. Mlynarski (1968) 
admitted that the bad condition of the material makes detailed descriptions largely 
a product of guesswork. 


Kansuchelys ovalis Yeh, 1963a:33 


Type.—IVPP V.1007, nearly complete shell. Type locality: Unknown (? Yushe, 
Shansi). Horizon: Unknown. 

Remarks.—Yeh (1963a) distinguished this species from K. chiayukuanensis on 
the basis of plastral scute and bone patterns. He further hypothesized that Kan- 
suchelys was more primitive than Testudo, but closely related. Auffenberg (1974) 
suggested that a specimen with such inexact data should not have been described. 

The entoplastron is overlapped by the gular scutes but not by the pectoral 
scutes. The supracaudal scute is entire and the anals are longer medially than the 
femorals. There are two suprapygals, the larger superior element embracing the 
inferior elliptical element. The neural pattern is 4-6-6-6-6-6-6-6, and the hexagonal 
neurals are short-sided anteriorly. 


Testudo shansiensis Wiman, 1930:38 


Type.—IVPP (Catalogue number not published), complete carapace, and plas- 
tron missing the anterior lobe. Type locality: southern part of locality 114, Shansi, 
Ho-Ch’u-Hsien, SE 60 li (1 li equals about one-third of a mile) of Nan-Skha-Wa, 
1 li from Ma-Hua-T’an. Horizon: Pliocene. 

Remarks.—Yeh (1963) referred two IVPP specimens to this species (IVPP 
V.1015, V.1016) and noted its similarity to T. hypercostata. Auffenberg (1974) 
allocated this form to the synonymy of T. hypercostata, which has the same type 
locality. 

The large dorsal suprapygal element is asymmetrical; only one ‘arm’ embraces 
the smaller elliptical ventral suprapygal. The anal and femoral scutes have similar 
median lengths. The gulars overlap the entoplastron. The pectoral scutes form a 
V-shaped sulcus which is posterior to the entoplastron on the midline but overlaps 
it laterally. This is similar to the condition in Testudo horsfieldii. The supracaudal 


VOLUME 96, NUMBER 3 573 


is entire. The neural formula is 4-8-4-8-4-6-6-6. Three marginal scutes contact the 
second pleural scutes. Whether a valid species or not, this form is certainly 
referable to Testudo. 


Testudo sharanensis Yeh, 1965:53 


Type.—IVPP V.2868, complete plastron, partial carapace missing some ele- 
ments of right side, and some fragments. A paratype was designated (IVPP 
V .2868.1) and consists of a complete plastron and a partial carapace. Type lo- 
cality: Ula Usu, Inner Mongolian Autonomous Region. Horizon: Late Eocene. 

Remarks.—Yeh (1965) considered this a very primitive species but discounted 
any close relationship to Hadrianus or Stylemys since all the neurals of these 
two are hexagonal whereas one neural (#2) of T. sharanensis is octagonal. He 
distinguished this species from Sinohadrianus sichuanensis on the basis of neural 
pattern differences. The position of the pleuro-marginal sulcus distinguished T. 
sharanensis from T. ulanensis Gilmore (i931), in which it extends along the lateral 
shell below the costo-peripheral suture. 

The gular scutes, but not the pectoral scutes overlap the entoplastron. There 
is plastral constriction at the lateral borders of the anal and gular scutes. The 
medial length of the anal exceeds that of the femoral scutes. The supracaudal 
scute is divided. Although Yeh characterizes the cervical as ‘‘small,’’ the cervical 
scute is large and broad in his fig. 3 (1965). There are notches in the anterior edge 
of the epiplastra. Yeh believes there are three suprapygal elements. I think it 
more likely that there is a supernumerary ninth neural yielding the following 
neural formula: 4-8-4-6-6-6-6-6-4. If this is the case, there are two suprapygal 
elements showing the primitive condition with the larger element embracing the 
more ventral elliptical element. These features suggest that this species is refer- 
able to ‘Manouria.’ 

Auffenberg (1974) did not refer to this species in his checklist. Chkikvadze 
(1973) contended that 7. sharanensis possesses intergular scutes; I disagree. Mly- 
narski (1976) referred this species to the Testudinidae incerta sedis and noted 
posterior plastral similarities shared with Ergilemys. 


Testudo shensiensis Wiman, 1930:28 


Type.—IVPP (Catalogue number not published), complete shell. Type locality: 
locality 51, Shensi, Fu-Ku-Hsien, W 110 li Wu-Lan-Kou, SE 1 li Pei-Hou-Kou. 
Horizon: Pliocene. 

Remarks.—Yeh (1963a) did not comment on the relationships of these species 
but refers four IVPP specimens to this taxon (IVPP V.1008, V.1009, V.10i0, 
V.1019). Glaessner (1933) allied this form to the antiqua-graeca phyletic line. 
Auffenberg’s (1974) cited Gilmore’s (1931) [actually 1934] assignment of AMNH 
material to this species. Gilmore was critical of Wiman’s (1930) work and pre- 
dicted that further study would reveal that fewer species should be recognized. 

The gular, but not pectoral scutes, overlap the entoplastron. The median length 
of the anal scutes is slightly less than that of the femoral scutes. There is no 
plastral constriction at the lateral border of either the gular or anal scutes. The 
cervical scute is relatively large according to Wiman’s (1930) figure (Pl. V, Fig. 
1). Three marginal scutes (numbers 5, 6 and 7) contact the second costal. The 


574 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


peey, 
ih 
pina 
r) 
r) @ é 
500 km o/¢ 


115 125 130 


Fig. 1. Localities in China and Mongolia where testudinid fossils have been unearthed (taken 
mostly from Yeh 1963a). The shaded area in the lower left is the present distribution of land tortoises 
in southeast Asia. 


neural formula is 4-8-4-6-6-6-6-6. There are two suprapygal elements of about 
equal size divided by a transverse suture. This species is best allocated to Tes- 
tudo. 


Sinohadrianus sichuanensis Ping, 1929:232 


Type.—Geological Survey of. China (Catalogue number not published), nearly 
complete shell. Type locality: ‘‘It was collected from Fan Chuan of Si Chuan 
Hsien, Honan Province by Mr. C. Li, .... The locality, where the specimen 
was secured, is near the border of Chieh Chia Chi of Chun Hsien, Hupei Province, 
where Mr. Li’s party happen to go across the boundary line between the two 
provinces while surveying’’ (Ping 1929:231). I have tried to locate this ambiguous 
type-locality without success. It is likely that this specimen was found north of 
the Han River on the northwestern border of Hupei Province and the south- 
western border of Honan Province in the Mot’ien Ling Mountains. Horizon: Late 
Eocene. 

Remarks.—Auffenberg (1974) suggested that Sinohadrianus is more primitive 
than North American Hadrianus and concurs with Yeh (1963a) that it is not 


VOLUME 96, NUMBER 3 575 


closely related to Hadrianus (=junior synonym of Manouria, according to Auf- 
fenberg 1971). 

The entoplastron is not overlapped by the pectoral scutes. Because the epi- 
plastra and the anterior entoplastron are missing, it is not possible to determine 
if the gular scutes overlap the entoplastron. Other plastral sutures are faint, but 
it seems that the median length of the anal and femoral scutes are near the same. 
Both the anterior and posterior margins of the carapace are missing, so it is 
impossible to determine the morphology of the cervical, suprapygal(s), and su- 
pracaudal(s). The neural formula is 4-6-6-6-4-6-6-6, and there may be a ninth 
neural whose shape is uncertain. 

Mlynarski (1976) included this species in the Testudinidae incerta sedis. If this 
species is a tortoise it might be referable to ‘“Manouria.’ 


Testudo sphaerica Wiman, 1930:33 


Type.—IVPP (Catalogue number not published), nearly complete carapace and 
plastron missing posterior lobe. Type locality: locality 110,, Shansi, Pao-Te-Chou, 
25 li NE of Chi-Chia-Kou, 5.5 li NE of Wang-Cia-Liang-Kou. Horizon: Pliocene. 

Remarks.—Mlynarski (1955) removed this form from the antiqua-graeca line 
claiming its affinities lie with 7. horsfieldii. He later (Mlynarski 1968) changed his 
mind and allocated this species to the Geochelone. Yeh (1963a) referred four 
IVPP specimens to this taxon (IVPP V.701, V.1011, V.1012, V.1013). Auffenberg 
(1974) followed Gilmore (1934) in considering 7. yushensis a synonym. Yeh 
(1963a) also referred the type of Terrapene sinica (IVPP V.701) and material 
unearthed from the Pliocene sediments of Yushe, Shansi (IVPP V.1011—V.1013) 
to this species. 

The gular, but not the pectoral scutes, overlap the entoplastron. Although the 
posterior lobe of the plastron is missing in the type, I expect that when discovered 
the anal scute will have about the same median length as the femoral scutes. 
There are two suprapygals of uncertain shape. The shape of the cervical scute is 
unknown. Only the anterior five neurals are known; the neural formula is 4-8-4- 
8-4?-?-?-?. Three marginal scutes contact the second pleural scute. Whether valid 
or not, this form is best allocated to Testudo. 


Kansuchelys tsiyuanensis Yeh, 1979:310 


Type.—IVPP (Catalogue number has not been published), complete shell. Type 
locality: Jiyuan, Henan Province. Horizon: Eocene. 

Remarks.—TYhis recently-named species is very primitive as Yeh (1979) noted. 
Neither the gular nor the pectoral scutes overlap the entoplastron. There is slight 
plastral constriction at the lateral anal border, but no constriction at the lateral 
gular border. The median length of the anal scutes is only slightly less than that 
of the femoral scutes. There are faint notches in the anterior lip of the epiplastral 
projection. The pectoral scutes narrow abruptly as they extend medially. The 
cervical is moderately broad dorsally, and may be even broader ventrally. There 
are two suprapygals; the larger superior element embraces the smaller ventral 
elliptical element. The supracaudal scute is divided. Most of the neurals are 
hexagonal with the short sides anterior; the neural formula is 4-6-6-6-6-6-6-6. The 
costal bones are alternatingly wide and narrow, medially and laterally. Marginals 


576 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


6, 7 and 8 contact the second pleural scute. This species is best allocated to 
‘Manouria.’ 


Testudo tungia Yeh, 1963b:224 


Type.—IVPP V.2768, slightly broken carapace. Type locality: Gigantopithecus 
cave, Liucheng, Kwangsi. Horizon: Early Pleistocene. 

Remarks.—Auffenberg (1974) believed this material is of a Cuora species. The 
neural formula, 4-6-4-4-4-4-?-?, supports this contention. Furthermore, the small 
size and characteristically emydid shell silhouette suggest that this species is not 
a testudinid. The plastron is not known. 


Testudo tunhuanensis Yeh, 1963a:42 


Type.—IVPP V.1029a, damaged anterior two-thirds of carapace. IVPP V.1029b, 
a complete plastron, is a paratype. Type locality: Taben-buluk, Tunhuang, Kansu. 
Horizon: Unknown (? Miocene). 

Remarks.—This species was described but not named by Bohlin (1953). Yeh 
(1963a), followed by Auffenberg (1974), noted the similarity between this form 
and T. sphaerica. 

The gular, but not the pectoral scutes, overlap the entoplastron. The median 
length of the anal scutes exceeds the median length of the femoral scutes. There 
is no plastral constriction at the lateral gular or anal borders. The cervical is 
moderate sized. The posterior half of the carapace is unknown. The neural for- 
mula is 4-8-4-7-5-6-6?-?. Probably the odd-sided neurals are in most specimens 
even-sided, as is usual in tortoises. Marginals 5, 6 and 7 contact the second pleural 
scute. 


Testudo ulanensis Gilmore, 1931:245 


Type.—AMNH 6691, a posterior plastral lobe attached to part of the left bridge, 
the median part of the plastron posterior of the entoplastron, a costal and pe- 
ripherals 9, 10 and 11 of both the right and left sides of the carapace with 
some fragments of costal bones articulated. Type locality: North Mesa, Shara 
Marun Region, Inner Mongolia. Horizon: Ulan Shireh, Upper Eocene. 

Remarks.—This difficult-to-allocate species was referred to Geochelone by 
Mlynarski (1968), and retained there with some reservation by Auffenberg (1974). 
Chkikvadze (1970), without stating his reasons, referred this species to Hadri- 
anus. 

The median length of the anal scutes is about equal to or exceeds the median 
length of the femoral scutes. The femoral scutes are separated from the inguinal 
scutes by the abdominal scutes. This condition is common in Testudo and ac- 
companies the development of a hinge in the posterior plastral lobe. There is 
plastral constriction at the lateral border of the anal scutes, but not at the lateral 
border of the gular scutes. 


Testudo yunnanensis Yeh, 1963a:47 


Type.—IVPP V.1031, portions of anterior plastron. Type locality: Wa-yao- 
chung, Ta-i-ma, Lunan. Yunnan. Horizon: Early Oligocene. 


VOLUME 96, NUMBER 3 Si 


Table 1.—The recommended nomenclatorial status of Chinese and Mongolian fossil tortoises. 


Original trivial name Original generic name Recommended generic name 
chiayukuanensis Kansuchelys “Manouria’ 
chienfutung ensis Testudo Testudo (?) 
demissa Testudo Geochelone 
hipparionum Testudo Testudo 
honanensis Testudo Testudo 
hyercostata Testudo Testudo 
insolitus Testudo Geochelone 
kalganensis Testudo Geochelone 
Kaiseni Testudo Indotestudo 
lunanensis Testudo Geochelone (?) 
nanus Testudo Indotestudo 
oskarkuhni Geochelone Geochelone 
ovalis Kansuchelys ‘Manouria’ 
shansiensis Testudo Testudo 
sharanensis Testudo “Manouria’ 
shensiensis Testudo Testudo 
sphaerica Testudo Testudo 
sichuanensis Sinohadrianus ‘“Manouria’ 
tsiyuanensis Kansuchelys “Manouria’ 
tungia Testudo Cuora 
tunhuanensis Testudo Testudo 
ulanensis Testudo Testude (?) 
yunnanensis Testudo Geochelone 
yushensis Testudo “Manouria’ (?) 


Remarks.—Auffenberg (1974) referred this species to Geochelone on the basis 
of size and believed T. /unanensis to be a junior synonym. Yeh (1963a) described 
the form because it is larger than most other Chinese tortoises. Chkikvadze (1972) 
allocated this species to his genus Ergilemys. 

The pectorals do not overlap the entoplastron and narrow abruptly as they 
extend medially. They are slightly broader on the midline than parasagittally. This 
condition of the pectoral scutes is also present in Geochelone sulcata and some 
Hesperotestudo species. The gular scutes overlap the entoplastron. The median 
length of the femoral scutes greatly exceeds the median length of the anal scutes. 
There is prominent constriction at the lateral borders of the anal scutes and the 
postero-lateral tips of the xiphiplastra are elongate and thickened. Most of the 
carapace is unknown. 


Testudo yushensis Yeh, 1963a:40 


Type.—IVPP V.1028, almost complete shell. Type locality: Tsuan-tse-yao, 
Yushe, Shansi. Horizon: Pliocene. 

Remarks.—Yeh (1963a) was careful to distinguish this species from T. hona- 
nensis but did so on the basis of highly variable characters. For this reason, 
Auffenberg (1974) considered this form a synonym of T. sphaerica. 

The gular, but not the pectoral scutes, overlap the entoplastron. The median 
length of the anal scutes is less than the median length of the femoral scutes. 
Nevertheless, the anal scutes are large and not reduced as in Geochelone. There 


578 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Table 2.—The stratigraphic position of the testudinids of China and Mongolia is summarized below. 
Exact stratigraphic information is, at present, not available for most Chinese sediments. Correlation 
with North American and/or European sediments has not yet been attempted. 


Epoch Species 
Pleistocene tungia* 
Pliocene hipparionum, hypercostata, oskarkuhni, shansiensis, 
shensiensis, sphaerica, yushensis 
Miocene chienfutungensis (?), honanensis (?), insolita, tunhuanensis (?) 
Oligocene—Late lunanensis, yunnanensis 
—Early demissa, kaiseni, nanus (?) 
Eocene—Late chiayukuanensis (?), sharanensis, sichuanensis, ulanensis 


—Middle or Early _—_ tsiywanensis 


Unknown horizon kalganensis, ovalis 


*_not a testudinid, probably Cuora (Auffenberg, 1974). 
(?)—stratigraphic position uncertain. 


is some plastral constriction at both the gular and anal lateral borders. The cer- 
vical scute is large and broadest posteriorly. There are two suprapygal elements; 
the larger superior element embraces the ventral elliptical element. The supra- 
caudal is entire. The neural formula is 4-6-4-8-4-6-6-6. This species may best be 
allocated to “Manouria.’ 


Discussion 


Although simple classifications have been advanced, there are no comprehen- 
sive phylogenetic hypotheses that incorporate Chinese and Mongolian fossil tor- 
toises. Hypotheses have been formulated by allocating species to particular gen- 
era; subsequent re-allocation, although reasonable, often did not include discussion 
of the data upon which the re-allocations rely (Auffenberg 1974). The above 
checklist constitutes the first step in a re-assessment of the phylogenetic rela- 
tionships of oriental fossil testudinids. However, this review relies extensively 
upon the literature and not upon a first hand examination of the material at present 
stored in the IVPP. For this reason, extensive changes in the classification of 
Chinese and Mongolian tortoises are deferred. 

Four genera have been recognized from eastern Asia: Testudo, Geochelone, 
Kansuchelys and Sinohadrianus. None of these are well diagnosed; character- 
izations prevail instead of comparative diagnoses. I recommend that four genera 
continue to be recognized, but different genera than those presently accepted. 
Diagnoses are below. 

Testudo: Testudo is the only testudinid (except Pyxis [Bour 1981] and Gopherus 
berlandieri [J. Howard Hutchison, pers. comm.] that develops an incipient hinge 
correlated with egg deposition in females) which normally possesses a posterior 
plastral hinge at the femoral-abdominal sulcus. Not all members of this genus 
have this hinge (e.g., Testudo horsfieldii) and it is sexually dimorphic in some 
species (Loveridge and Williams 1957). Tortoises of this genus differ further from 
Geochelone in possessing enlarged anal scutes whose median length is equal or 
exceeds that of the femoral scutes. Testudo differs from ‘Manouria’ by possessing 


VOLUME 96, NUMBER 3 579 


an entire supracaudal scute. Testudo is distinguished from both of the above 
genera in having undifferentiated marginals in which three marginals contact the 
second pleural scute (Roger Bour, pers. comm.). In Testudo, unlike Indotestudo, 
the pectoral scutes do not overlap the entoplastron; or if such overlap occurs it is 
parasagittal rather than medial. 

Geochelone: Geochelone differs from all other Chinese tortoises except In- 
dotestudo, by possessing small anal scutes whose median length is very much 
less than the median length of the femoral scutes. Unlike Indotestudo, the pec- 
toral scutes do not overlap the entoplastron. 

Indotestudo: This genus differs from all other tortoise genera by possessing a 
transversely extending humeral-pectoral sulcus which crosses the entoplastron. 
Bour’s (1980) elevation of this subgenus of Geochelone to generic level is fol- 
lowed. 

‘Manouria’: This primitive genus may or may not be monophyletic. It differs 
from all other tortoises in having a divided suprapygal which is always divided 
on both its dorsal and ventral surfaces. (In Malacochersus and T. hermanni the 
supracaudal is entire ventrally and divided dorsally.) Furthermore, unlike most 
tortoises, except Stylemys, the neurals are hexagonal and tetragonal, and usually 
not octagonal. Recent “Manouria’ (i.e., Geochelone emys and G. impressa), 
sometimes have octagonal neurals anteriorly. Also unlike most other tortoises, 
except Malacochersus, the epiplastral projection is not greatly thickened, but is 
flattened and often concave upon its dorsal surface. Hadrianus is included within 
‘Manouria’ as suggested by Auffenberg (1971). Kansuchelys and Sinohadrianus 
are here referred to “Manouria.’ ‘Manouria,’ although easily recognizable, is 
presently diagnosed by shared primitive features. 


Acknowledgments 


I wish to thank Gene Gaffney, George Zug, Carl Ernst, Sam McDowell, Rich- 
ard Vari, and Yeh Hsiang-k’uei for their help during the preparation of this check- 
list. The comments of an anonymous reviewer increased the clarity of this paper. 
This work was completed during my tenure as a predoctoral fellow at the National 
Museum of Natural History, Smithsonian Institution. 


Literature Cited 


Andrews, C. W. 1906. A descriptive catalogue of the Tertiary Vertebrata of the Fayum, Egypt.— 
British Museum (Natural History), 324 pp. 

Auffenberg, W. 1962. A new species of Geochelone from the Pleistocene of Texas.—Copeia 1962(3): 
627-636. 

——. 1963. Fossil Testudinine turtles of Florida: Genera Geochelone and Floridemys.—Bulletin 
of the Florida State Museum 7(2):53—97. 

———. 1966. The carpus of land tortoises (Testunidae).—Bulletin of the Florida State Museum 
10(5): 159-191. 

—. 1971. A new fossil tortoise with remarks on the origin of South American tortoises.— 
Copeia 1971(1):106—117. 

—. 1974. Checklist of fossil land tortoises (Testudinidae).—Bulletin of the Florida State Mu- 
seum 18(3):121—251. 

— —. 1976. The genus Gopherus (Testudinidae). Pt. I. Osteology and relationships of extant 
species.—Bulletin of the Florida State Museum 20(2):47-120. 

Bohlin, B. 1953. Fossil reptiles from Mongolia and Kansu.—Reports of Scientific Expedition, 


580 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Northwestern Provinces of China. VI. Vertebrate Paleontology, no. 6. Statens Ethnografiska 
Museum (Stockholm), 109 pp. 

Bramble, D. M. 1971. Functional morphology, evolution, and paleoecology of Gopher tortoises.— 
Unpub. Ph.D. Dissertation, University of California, Berkeley, 351 pp. 

—. 1982. Scaptochelys: Generic revision and evolution of Gopher tortoises.—Copeia 1982(4): 
852-867. 

de Broin, F. 1977. Contribution a L’etude des Cheloniens. Cheloniens continentaux du Cretace et 
du Tertiaire de France—Memoires du Muséum National d’Histoire Naturelle ser. C, 38:1— 
366, 115 figs., 38 pls. 

Bour, R. 1981. Etude systematique du genere endemique Malagache Pyxis Bell, 1827 (Reptilia, 
Chelonii).—Bulletin de Société Linneene de Lyon 50(4): 132-176. 

—. 1980. Essai sur la taxonomie Testudinidae actuels (Reptilia, Chelonii).—Bulletin de Muséum 
National d’ Histoire Naturelle, Paris 4, 2, A(2):541-546. 

Chkikvadze, V. 1970. Klassifikacija podklasa Testudinat.—l6aja Naucnaja sesia Institute Paleo- 
biologii AN Gruziya SSR., pp. 7-8, Tbilisi. 

——. 1972. [On the systematic position of Tertiary gigantic land tortoises of the Palearctic].— 

Bulletin Academy of Sciences, Georgian SSR 65(3):745-748. [In Russian. ] 

1973. Treticnyje cerepachi Zajsanskoj Kotliviny.—°*Macnierba,”’ 100 pp., Tbilisi. 

Gilmore, C. W. 1931. Fossil turtles of Mongolia.—Bulletin of the American Museum of Natural 
History 59:213-257. 

—. 1934. Fossil turtles of Mongolia: A second contribution.—American Museum Novitates 
689: 1-14. 

Glaessner, M. F. 1933. Die Tertiarschildkroten Niedrosterichs.—Neues Jarhbuch ftir Minerologie, 
Geologie und Paleontologie, Abteilung B 69:353—387. 

———. 1935. Bemerkungen zur tertiaren Schildkrotenfauna.—Ungarisches Zentralblatt fiir Mine- 
rologie, Abteilung B, pp. 124-129. 

Gradzinski, R., J. Kazmierczak, and J. Lefeld. 1968. Geographical and geological data from the 
Polish-Mongolian Palaeontological Expeditions. Results of the Polish-Mongolian Palaeontolog- 
ical Expeditions, I.—Palaeontologia Polonica 19:33-82. 

Hutchison, J. H. 1980. Turtle stratigraphy of the Willwood formation, Wyoming: Preliminary re- 
sults. Jn Early Cenozoic Paleontology and Stratigraphy of the Bighorn Basin, Wyoming.— 
University of Michigan, Papers in Paleontology 24:115-118. 

Khozatsky, L. E. 1953. Voprosy Paleontologii 1:20-31. 

Loveridge, A., and E. E. Williams. 1957. Revision of the African tortoises and turtles of the 
suborder Cryptodira.—Bulletin of the Museum of Comparative Zoology, Harvard 115:163—557. 

Matthew, W. D., and W. Granger. 1923. The fauna of the Ardyn Obo formation.—American Mu- 
seum Novitates 98:1—5. 

Mlynarski, M. 1955. Zolwie zu Pliocenu Polski.—Acta Geologica Polonica 5(2):161—214. 

1968. Results of the Polish-Mongolian Paleontological Expedition. Pt. I. Notes of the tor- 

toises of the Tertiary of Mongolia.—Palaeontologia Polonica (19):85—97. 

1976. Testudines.—Handbuch der Palaoherpetologie 7: 1-130. 

Ping, C. 1929. A new Eocene land turtle from Honan.—Bulletin of the Geological Society of China 
8(3):23 1-238. 

Schleich, H.-H. 1981. Jungtertiare Schildkroten Suddeutschlands unter besonderer Berucksichti- 
gung der Fundstelle Sandelhausen.—Courier Forschungsinstitut Senckenberg 48: 1-372. 

Van Devender, T., K. B. Moody, and A. H. Harris. 1979. The desert tortoise (Gopherus agassizii) 
in the Pleistocene of the northern Chihuahuan desert.—Herpetologica 32:298—304. 

Wiman, C. 1930. Fossile Schildkroten aus China.—Paleontologica Sinica (Peking) ser. C 6(3):1—57. 

Yeh, H.-k. 1963a. Fossil turtles of China.—Palaeontologica Sinica (150): 1-122. 

1963b. A new Quaternary Testudo from the Gigantopithecus cave, Liucheng, Kwangsi.— 

Vertebrata PalAsiatica 7(3):224—229. 

——. 1965. New materials of Fossil turtles of Inner Mongolia.—Vertebrata PalAsiatica 9(1): 
48-69. 

—. 1979. Fossil testudinids from Jiyuan, Henan.—Vertebrata PalAsiatica 17(4):310-317. 


Division of Amphibians and Reptiles, Department of Vertebrate Zoology, Na- 
tional Museum of Natural History, Smithsonian Institution, Washington, D.C. 
20560. 


PROC. BIOL. SOC. WASH. 
96(3), 1983, pp. 581-597 


NEOTROPICAL MONOGENEA.* 5. FIVE NEW SPECIES 
FROM THE ARUANA, OSTEOGLOSSUM BICIRROSUM 
VANDELLI, A FRESHWATER TELEOST FROM BRAZIL, 
WITH THE PROPOSAL OF GONOCLEITHRUM N. GEN. 
(DACTYLOGYRIDAE: ANCYROCEPHALINAE) 


Delane C. Kritsky and Vernon E. Thatcher 


Abstract.—Gonocleithrum n. gen. (Dactylogyridae: Ancyrocephalinae) is pro- 
posed for five new species collected from the gills of the aruana, Osteoglossum 
bicirrosum Vandelli, from Janauaca Lake, near the south bank of the Solimoes 
River, Manaus, Amazonas, Brazil, as follows: G. planacrus, G. aruanae, G. 
coenoideum, G. cursitans, and G. planacroideum new species. Gonocleithrum is 
similar to Urocleidoides Mizelle and Price, 1964, but has a ventral gonadal bar 
lying near the anterior end of the ovary. A table is presented showing the known 
freshwater monogenean fauna of fishes from the Neotropical Region. 


The Neotropical Region and particularly South America support an unique 
freshwater monogenean fauna which differs significantly from that of North 
America. Of the 26 known genera of Monogenea from this region (Gonocleithrum 
n. gen. included), 23 are at present restricted under natural conditions to the 
neotropics (Table 1). In addition, re-examination of the eight species included in 
Cleidodiscus and Urocleidus (both North American genera) will undoubtedly 
result in their transfer to other genera unique to the Neotropical Region. 

Gussev (1978) suggests that the South American fauna of Monogenea has an 
ancient evolutionary relationship to that of Africa and in fact considers the Af- 
rican genus Characidotrema a junior synonym of Jainus. While we do not accept 
this synonymy, since it is based on information published in original articles and 
not on reexamination of the species involved, the apparent resemblance between 
species of these genera provides support for Gussev’s hypothesis. Although studies 
on the Neotropical and the Ethiopian (African) Monogenea have just begun to de- 
termine the faunas present, a more thorough knowledge of both is necessary to 
substantiate the relationship. 


Material and Methods 


The host, Osteoglossum bicirrosum Vandelli, was collected on two occasions 
from Janauaca Lake, near the south bank of the Solimoes River, Manaus, Ama- 
zonas, Brazil, on 21 March and 14 August 1978, respectively. These fish were 


* The title of our series on Neotropical monogeneans is changed to conform with resolutions of 
the Round Table Discussion, Monogenea: Problems of Systematics Biology and Ecology, held on 23 
August, 1978 during the IV International Congress of Parasitology in Warsaw, Poland (Euzet and 
Prost 1981, Review Advances Parasitology, Warszawa 1003-1004). 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


582 


(PL61) JoyoIeYy, pure AysI Ly 
(pL6l) JoyoyveyL pue Asi 
(6961) O19WOY pue dd1Ig 


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poiquvssow pidoly 
‘ds nuisypjdopnasg 


avid “Ds 
SHSOAI]IS ‘Ox 


sisuadvnsv.ivd “vy 


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WiN1YpajIOYdu y 
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(6L6]) UoIAR yy pue ‘1oYyo ey, ‘AYSIU sniajdouvjou Uuookig SNAMIO[DAIAS “VW 
(6L6[) UoARy pue ‘19Yyoeyy, ‘AYSILy uUnNuUodosIDU *D 
(6L6|) UoARy pues ‘1ayoyeuy, ‘AYSIU suaplg DUWOSSO]OD snypinyivds “vy 
(S961) 99g pure s][ezI/\ MAdAIJIDU SNUJVSSDAIIS syvoido1joau “Vv 
(6L6[) UoARyY pue ‘194s ey, ‘AYSILLy] Sniajdouvjaw UuodKig Mamapimniy “VW 
(6L6[) UoWARy pue ‘1ayoeYyy, ‘AYSIL sniajdouvjow UuodKig SuDda]a “VW 
(PL61) JoyoIeU, pue AXSILWyy] siuyffp snumjos SNUIGDAOJNIYNI “VW 
(PL61) oydIVYL pue AYys}LUy siuyffp snunupos SNUDIQUO]OI “VY 
(46961) AXSHIM pure sI[IZIA Snsajdouvjaw UOdKAg SIAIIG “VW 
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(S961) POU pur sI[OZIIN MIAIPIDU SNUJVSVAII SNAOYJUDIVUD “VY snsioyjUuvovUuy 
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(sigou) Jayoyeyy, pue AXsLy WINSOAMIIG UNSSO]S0IISE uinaplosovunjd *H) 
(Sigou) Jayoyeyy, pue AYsLTy WINSOAMIIG UNSSO]SO0I1SE SUDJISAND “D 
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(F681) Sopuvig MUOSIIWIWOD SNUY DJODIAO * Tue. DIJAUDIDIAAP M7 
(L961) UNMON pue sug sp3is puivdviy WINLJSIDUDOJIKI “G DUIAJSIMDG 
(S961) 99Lg pue s][ezIpy 11949]JDU SNUIDSDASOS SNUIDSDAIAS “Dd 
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VOLUME 96, NUMBER 3 


ee 
goualajoy JsoH auseied 
Neen eee ee EEE 


‘panunuog— | 21921 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


584 


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(86961) AYSIU pure s[[ezIW ‘ds pipwvyy snjsngos “f) 
(p961) 99 pue s|[IZIP DIVINIYIA SI1S1GIT SNIDINIYIA “(| 
(796) Buledeg pue uyoy DIDINIYAdA SAJSIGaT DINUIW * se sexe 
(996]) OUBID pue AYSILY “S[[IZIP SNWUOISOADIU SNUMIDAS U9 FT SNUOJSOADIU * (1) 
(86961) AYSIY pure o[fezIp\ DUI] WIgnsosy SIYIIOSIU *() 
(PL61) Opueuloy pue ‘youry ‘reujoyy Snauav SVAOPKIOD ISOSADU *D 
(9L61) 1o4uDIeYL pue AYs}Ly o1nsunz sninjisojpyday 1AaDUIDUI * 
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‘panunuog—'| 21921 


585 


VOLUME 96, NUMBER 3 


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ddUd1BJOY JsOoH oyiseieg 


‘penunuoy— | 219e,L 


586 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


treated and parasites collected from the gills and stored according to the proce- 
dures of Kritsky and Thatcher (1974). Parasites were stained with Mayer’s car- 
mine or Gomori’s trichrome and mounted in permount for observing internal 
organs. Other specimens were mounted unstained in Gray and Wess’ medium for 
study of sclerotized structures. Measurements of parasites were made according 
to the guidelines of Mizelle and Klucka (1953) except that the cirrus measurement 
represents the diameter of the first ring (proximal) of the cirrus coil; all measure- 
ments are in micrometers. Figures were prepared with the aid of a microprojector 
or camera lucida. Type-specimens are deposited in the collections of the Instituto 
Nacional de Pesquisas da Amazonia (INPA), the helminthological collection of 
the National Museum of Natural History, Smithsonian Institution (USNM), and 
the University of Nebraska State Museum (UNSM) as indicated below. 


Gonocleithrum, new genus 


Diagnosis.—Dactylogyridae, Ancyrocephalinae. Body divisible into cephalic 
region, trunk, peduncle, and haptor. Tegument thin, smooth. Cephalic lobes, head 
organs, cephalic glands present. Four eyes. Mouth subterminal, midventral; phar- 
ynx muscular, glandular; esophagus present; intestinal crura 2, confluent poste- 
rior to testis, lacking diverticulae. Gonads intercecal, overlapping; testis dorso- 
posterior to ovary. Vas deferens looping left intestinal crus; seminal vesicle a 
simple dilation of vas deferens; copulatory complex comprising coiled or modified 
coiled cirrus, accessory piece. Vagina sinistral, seminal receptacle present. Ven- 
tral Y-shaped gonadal bar lying near anterior end of ovary. Vitellaria well de- 
veloped. Haptor armed with dorsal and ventral pair of anchors, dorsal and ventral 
bar, 7 pairs of flexible hooks with ancyrocephaline distribution (Mizelle 1936). 
Parasites of Osteoglossidae. 

Type-species and host.—Gonocleithrum planacrus n. sp. from Osteoglossum 
bicirrosum Vandelli, Janauaca Lake, near the south bank of the Solimoes River, 
Manaus, Amazonas, Brazil. 

Other species.—Gonocleithrum aruanae n. sp., G. coenoideum n. sp., G. cur- 
sitans n. sp., G. planacroideum n. sp., all from Osteoglossum bicirrosum. 

Remarks.—Gonocleithrum is similar to the Neotropical genus Urocleidoides 
Mizelle and Price, 1964, in that the cirrus is comprised of a coil with few to many 
rings and by the general arrangement of the haptoral armament. The new genus 
differs from Urocleidoides by possessing a Y-shaped gonadal bar near the anterior 
end of the ovary. 

Based on the fact that Urocleidoides species occur on fishes from several orders 
and families, Gussev (1978) suggests that this genus may be an assemblage of 
species representing several different genera. In fact, if it were not for the pres- 
ence of the gonadal bar, the five following species could be placed in Uroclei- 
doides as it is defined at present. All five species of Gonocleithrum are from 
Osteoglossum bicirrosum which is a member of the primitive order of bony- 
tongued fishes, Osteoglossiformes. No species of Urocleidoides has been re- 
ported from this host group. 

Price and Nowlin (1967) reported Dawestrema cycloancistrium from the gills 
of Arapaima gigas (Osteoglossidae, Arapaiminae) in the Amazon River and its 
tributaries. Also, Paperna (1969) described Heterotesia voltae from Heterotis 


VOLUME 96, NUMBER 3 587 


niloticus (Osteoglossidae, Heterotinae) in Africa. Species of Gonocleithrum are 
distinguished from these ancyrocephalines infesting fishes of the Osteoglossi- 
formes by possessing a gonadal bar. 

The function of the gonadal bar is not clear, but it could be involved in ori- 
entation during copulation. The protruding anterior arms of the bar form a pouch- 
like structure on the ventral surface of the worm that could assist in positioning 
of the copulating partner. We feel that the gonadal bar is not analogous to the 
vaginal sclerite of some Urocleidoides species (U. reticulatus Mizelle and Price, 
1964, and U. anops Kritsky and Thatcher, 1974), which apparently functions as 
a supporting structure of the vagina. 


Gonocleithrum planacrus, new species 
Figs. 1-8 


Type-specimens.—Holotype, INPA-234-1; paratype, USNM 77377; paratype, 
UNSM 21480. 

Description (based on 3 specimens).—Body fusiform; length 616 (604—628), 
greatest trunk width 100 (98-102) near midlength. Cephalic lobes well developed, 
2 terminal, 2 bilateral; head organs large, lying in cephalic lobes and adjacent 
cephalic area; cephalic glands unicellular, situated in 2 bilateral groups postero- 
lateral to pharynx near ventral surface. Members of anterior pair of eyes smaller, 
usually closer together than members of posterior pair; eye granules small, irreg- 
ular to subovate; accessory granules absent or very few in proximity of eyes. 
Pharynx spherical, 35 (33-37) in diameter. Peduncle broad; haptor subhexagonal, 
147 (143-151) wide, 136 (128-144) long. Anchors similar in shape; each with large 
base, short shaft, point with subterminal bends; ventral anchor 83 (79-85) long, 
base 42 (35-50) wide; dorsal anchor 61 (60—64) long, base 38 (33-43) wide. Anchor 
filament variable, double. Ventral bar 71 (68-76) long, broad, rod-shaped, with 
anteriorly directed medial process; dorsal bar 80 (74-89) long, rod-shaped, with 
slight medial bend. Hooks similar; each with inflated proximal shank, depressed 
thumb, fine point; hook pairs 1, 2, 3, 4, 6, 7—31 (30-32) long, pair 5—20 to 21 
long; FH loop 4% shank length. Cirrus a coiled, heavily sclerotized tube, with 2 
rings, subterminal flange, finely tapered tip; diameter of complete ring 28 (27-29). 
Accessory piece a variable fleshy structure basally articulated to cirrus base. 
Testis subspherical, 38 to 39 in diameter; seminal vesicle inconspicuous; prostatic 
reservoirs 2, with thick walls; prostate a large crescent of cells located anterior 
to vitelline commissure. Ovary elongate ovate, 45 (42-49) wide, 105 (100-110) 
long; seminar receptacle, oviduct, ootype, uterus, genital pore not observed; 
Vagina an irregular and lightly sclerotized tube; vitellaria dense, coextensive with 
gut. Gonadal bar 62 to 63 long; anterior arms expanded, recurved. 

Remarks.—Gonocleithrum planacrus is the type-species for the genus. The 
specific name is from Greek (plano = wandering + acrus = tip) and refers to the 
shape of the anchor points. 


Gonocleithrum aruanae, new species 
Figs. 9-17 
Type-specimens.—Holotype, INPA-236-1; paratypes, INPA-236-2 to 5; para- 
types, USNM 77379; paratype, USNM 21482. 


588 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Figs. 1-8. Gonocleithrum planacrus: 1, Ventral view of holotype; 2, Gonadal bar; 3, Copulatory 
complex; 4, Hook; 5, Ventral anchor; 6, Dorsal anchor; 7, Ventral bar; 8, Dorsal bar. All figures are 
drawn to the same scale (30 micrometers) except Fig. 1 (100 micrometers). 


Description (based on 20 specimens).—Body foliform; length 334 (309-378), 
greatest trunk width 70 (52-91) in anterior half. Two terminal, 2 bilateral cephalic 
lobes inconspicuous; well-developed head organs usually in 4 distinct pairs; ce- 
phalic glands obscured by vitellaria. Eyes equidistant, members of anterior pair 


VOLUME 96, NUMBER 3 589 


12 : 
Ze pa \ 
14 
15 
17 


16 


Figs. 9-17. Gonocleithrum aruanae: 9, Composite drawing of whole mount (ventral); 10, Gonadal 
bar; 11, Copulatory complex; 12, Vagina and gonadal bar; 13, Dorsal bar; 14, Hooks; 15, Ventral 
bar; 16, Dorsal anchor; 17, Ventral anchor. All figures are drawn to the same scale (30 micrometers) 
except Fig. 9 (100 micrometers). 


smaller than those of posterior pair; eye granules ovate, medium in size; acces- 
sory granules throughout cephalic area. Pharynx spherical, 18 (15—20) in diameter. 
Peduncle moderately broad; haptor subhexagonal, 92 (68-106) wide, 71 (60-76) 
long. Anchors dissimilar; ventral anchor 45 (38-50) long, with well-developed 


590 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


roots, elongate shaft, sharply recurved point, base 20 (18-22) wide; dorsal anchor 
36 (34-38) long, with small base, curved shaft, straight point, base 14 (12-16) 
wide. Anchor filament variable, double. Ventral bar 36 (32-39) long, with enlarged 
terminations, anteriorly directed medial process; dorsal bar V-shaped, 35 (32-37) 
long. Hook pairs 1, 2, 3, 4, 6, 7—28 (24-32) long, similar; each with inflated 
shank, depressed thumb, fine point. Hook pair 5—15 (14-16) long, with basal 
inflation of shank, erect thumb, fine point. FH loop 4 shank length except pair 
5 (4% shank length). Cirrus a coil of about 7 rings, basal ring diameter 9 (8—10); 
accessory piece variable, lying within cirrus rings, articulated to cirrus base. 
Testis subspherical, 22 (15-27) in diameter; seminar vesicle poorly defined; pros- 
tatic reservoirs 2, each with thick wall; prostate not observed. Ovary elongate 
ovate 24 (20-31) wide, 53 (46-63) long; seminar receptacle, oviduct, ootype, uter- 
us, genital pore not observed; vagina a simple sclerotized tube with distal flare; 
vitellaria dense, coextensive with gut. Gonadal bar 30 (28-33) long, with anterior 
arms slightly longer than base; small truncate process near base of anterior arms. 

Remarks.—This species is easily confused with Gonocleithrum cursitans n. 
sp., with which it was found on Osteoglossum bicirrosum in about equal numbers. 
The anchors, bars, and hooks are nearly impossible to distinguish in these species. 
However, G. aruanae n. sp. is separated from G. cursitans by having 1) a stout, 
more robust gonadal bar, 2) cirrus rings with noticeably smaller diameter, and 3) 
a simple tubular vagina which lacks the proximal coils of G. cursitans. The species 
name is derived from the local name of the host. 


Gonocleithrum coenoideum, new species 
Figs. 18-25 


Type specimens.—Holotype, INPA-238-1; paratypes, INPA-238-2 to 6; para- 
types, USNM 77381; paratype, UNSM 21484. 

Description (based on 34 specimens).—Body fusiform; length 389 (294-477), 
greatest trunk width 80 (45-117) near midlength. Cephalic lobes inconspicuous, 
2 terminal, 2 bilateral; head organs well developed, one lying in each cephalic 
lobe and adjacent cephalic area; cephalic glands numerous, unicellular, lying 
posterolateral to pharynx. Eyes large; members of posterior pair larger, closer 
together than those of anterior pair; eye granules usually small, subovate; acces- 
sory granules usually absent. Pharynx spherical, 21 (18-23) in diameter. Peduncle 
broad; haptor subhexagonal, 67 (45-83) wide, 55 (38-76) long. Anchors similar, 
each with elongate superficial root, short shaft, long curved point; ventral anchor 
32 (28-34) long, base 16 (13-19) wide; dorsal anchor 31 (28-35) long, base 16 (13- 
18) wide. Anchor filament variable, double. Bars similar, broadly V-shaped; ven- 
tral bar 37 (32-42) long; dorsal bar 29 (25-34) long. Hooks similar, each with 
inflated proximal shank, depressed thumb, fine point; hook pairs 1, 2, 3, 4, 6, 7— 
22 (20-24) long, pair 5—19 (18-20) long. FH loop % shank length. Cirrus a coil 
with 3-4 rings, basal ring diameter 11 (10-12); accessory piece variable, basally 
articulated to cirrus base. Testis subovate, 12 to 13 in diameter; seminal vesicle 
large; prostatic reservoirs not observed. Ovary bacilliform, 18 (13-22) wide, 59 
(49-69) long; oviduct, ootype, uterus, genital pore not observed; vagina nonscler- 
otized; seminar receptacle subspherical, lying dorsal to gonadal bar; vitellaria 


VOLUME 96, NUMBER 3 59] 


23 


24 


30p 


, tie aie ohare 


‘S, af 


Figs. 18-25. Gonocleithrum coenoideum: 18, Composite drawing of whole mount (ventral); 19, 
Gonadal bar; 20, Copulatory complex; 21, Ventral anchor; 22, Dorsal anchor; 23, Hook; 24, Ventral 
bar; 25, Dorsal bar. All figures are reproduced to the same scale (30 micrometers) except Fig. 18 (100 
micrometers). 


592 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


dense, coextensive with gut. Gonadal bar 23 (20-26) long, with tapered arms and 
bulbous base. 

Remarks.—Gonocleithrum coenoideum is not closely related to any of the 
species in the genus. However, the structure of the copulatory complex suggests 
affinity to G. cursitans and G. aruanae. The specific name is from Greek (coeno = 
common + oides = like). 


Gonocleithrum cursitans, new species 
Figs. 26—33 


Type-specimens.—Holotype, INPA-237-1; paratypes, INPA-237-2 to 4; para- 
types, USNM 77380; paratype, UNSM 21483. 

Description (based on 18 specimens).—Body foliform; length 338 (279-378), 
greatest trunk width 69 (53-83) in anterior half. Two terminal, 2 bilateral cephalic 
lobes poorly developed; head organs well developed, 3 pairs; cephalic glands 
obscured by vitellaria. Eyes equidistant, members of posterior pair larger than 
those of anterior pair; eye granules elongate ovate, medium in size; accessory 
granules throughout cephalic region. Pharynx spherical, 15 (12-17) in diameter. 
Peduncle broad; haptor subhexagonal, 91 (75—121) wide, 66 (53-72) long. Anchors 
dissimilar; ventral anchor 43 (41-45) long, with well-developed roots, bent shaft, 
sharply recurved point, base 21 (20—23) wide; dorsal anchor 34 (32—36) long, with 
elongate superficial root, curved shaft, long point, base 14 (13-15) wide. Anchor 
filament variable, double. Ventral bar 40 (36-47) long, with enlarged ends, median 
anterior process; dorsal bar 40 (35-47) long, broadly V-shaped. Hook pairs 1, 2, 
3, 4, 6, 7 similar, each with inflated shank, depressed thumb, fine point; hook 
pair 1—36, pair 2—26 (25-27), pairs 3, 4, 6, 7—30 (28-33) long; Hook pair 5—15 
to 16 long, with inflated proximal shank, erect thumb, fine point. FH loop 4% 
shank length except hook 5 (% shank length). Cirrus a coil with 5—6 rings, basal 
ring diameter 20 (18-23); accessory piece a spiral rod lying within cirrus rings, 
basally articulated to cirrus base. Testis subspherical, 19 (15—23) in diameter; 
seminal vesicle poorly defined; prostatic reservoirs 2, with conspicuous walls. 
Ovary bacilliform, 23 (22—24) wide, 60 (48-72) long; oviduct, seminal receptacle, 
ootype, uterus, genital pore not observed. Vagina a delicate sclerotized tube, 
coiled anterior to gonadal bar; vitellaria dense, coextensive with gut. Gonadal 
bar with inconspicuous flanges on anterior arms, small medial truncate process 
near base of anterior arms; bar 30 (27-33) long. 

Remarks.—Gonocleithrum cursitans is obviously a close relative of G. arua- 
nae. Distinguishing characteristics, which include the morphology of the gonadal 
bar, copulatory complex, and vagina, are explained in the remarks for G. arua- 
nae. The specific name is from Latin (cursitans = running about). 


Gonocleithrum planacroideum, new species 
Figs. 34-41 


Type-specimens.—Holotype, INPA-235-1; paratypes, USNM 77378; paratype, 
UNSM 21481; INPA-235-2. 

Description (based on 6 specimens).—Body fusiform; length 372 (340-408), 
greatest trunk width 58 (53-72) at level of gonads. Two terminal, 2 bilateral ce- 
phalic lobes poorly developed; head organs well developed, usually 2 pairs; ce- 
phalic glands inconspicuous. Members of posterior pair of eyes larger, farther 


VOLUME 96, NUMBER 3 593 


Figs. 26-33. Gonocleithrum cursitans: 26, Composite drawing of whole mount (ventral); 27, Go- 
nadal bar; 28, Copulatory complex; 29, Ventral anchor; 30, Dorsal anchor; 31, Hooks; 32, Ventral 
bar; 33, Dorsal bar. All figures are drawn to the same scale (30 micrometers) except Fig. 26 (100 
micrometers). 


594 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


38 


30 


Figs. 34-41. Gonocleithrum planacroideum: 34, Ventral view of holotype (specimen slightly rolled 
to left); 35, Gonadal bar; 36, Copulatory complex; 37, Ventral anchor; 38, Dorsal anchor; 39, Hook; 
40, Ventral bar; 41, Dorsal bar. All figures are drawn to the same scale (30 micrometers) except Fig. 
34 (100 micrometers). 


VOLUME 96, NUMBER 3 595 


apart than those of anterior pair; eye granules irregular; accessory granules ab- 
sent. Pharynx spherical, 21 (19-22) in diameter; gut inconspicuous. Peduncle 
broad; haptor subhexagonal, 69 (64-76) wide, 64 (56-68) long. Anchors dissimilar; 
ventral anchor 41 (36-47) long, with large base, curved shaft, point with subter- 
minal bends, base 20 (15-23) wide; dorsal anchor 28 (25-30) long, with well- 
developed roots, curved shaft and point, base 12 (11-13) wide. Anchor filament 
not observed. Ventral bar 51 (46-58) long, rod-shaped, with expanded ends, me- 
dial anterior process; dorsal bar broadly V-shaped, 30 (29-31) long. Hooks sim- 
ilar, each with inflated proximal shank, erect thumb, fine point; hook pairs 1, 2, 
3, 4, 6, 7—23 (21-26) long, pair 5—17 to 18 long; FH loop 2/5 shank length. Cirrus 
a coiled tube with 3-4 rings, basal ring diameter 6 (5—7); accessory piece variable, 
not articulated to cirrus base. Testis subovate, 24 (16—32) in diameter; vas def- 
erens not observed; seminal vesicle large; prostatic reservoirs with delicate wall 
(one observed). Ovary bacilliform, 28 wide, 66 (68-70) long; oviduct, uterus, 
ootype, genital pore not observed; vagina tubular with variable diameter; seminal 
receptacle subspherical, lying dorsal to gonadal bar; vitelleria dense, coextensive 
with gut. Gonadal bar 25 (23-28) long, with enlarged terminations of anterior 
arms. 

Remarks.—This species most closely resembles Gonocleithrum planacrus as 
shown by the morphology of the ventral anchor. The two species are easily 
separated by the comparative morphology of the dorsal anchor, dorsal bar, go- 
nadal bar, and copulatory complex. The specific name, from Greek, indicates the 
relationship of these two species. 


Literature Cited 


Brandes, G. 1894. Fridericianella ovicola n. g., n. sp. Ein neuer monogenetishcher Trematod.— 
Abhandlungen der Naturforschenden Gesellschaft zu Halle Bd. XX. Jubilaums—Festschnift: 
303-310. 

Gussev, A. V. 1978. Monogenoidea of freshwater fishes. Principles of systematics, analysis of world 
fauna and its evolution.—Parazitologicheskii Sbornik 28:96—198. 

Hanek, G., K. Molnar, and C. H. Fernando. 1974. Three new genera of Dactylogyridae (Mono- 
genea) from freshwater fishes of Trinidad.—Journal of Parasitology 60:911-913. 

Kohn, A., andI. Paperna. 1964. Monogenetic trematodes from aquarium fishes.—Revista Brasileira 
de Biologia 24:145-149. 

Knitsky, D. C., and T. H. Fritts. 1970. Monogenetic trematodes from Costa Rica, with the proposal 

of Anacanthocotyle gen. n. (Gyrodactylidae: Isancistrinae).—Proceedings of the Helmintho- 

logical Society of Washington 37:63—68. 

, and P. D. Leiby. 1972. Dactylogyridae (Monogenea) from the freshwater fish, Astyanax 

fasciatus (Cuvier), in Costa Rica, with descriptions of Jainus hexops sp. n., Urocleidoides 

costaricensis, and U. heteroancistrium combs. n.—Proceedings of the Helminthological So- 

ciety of Washington 39:227-230. 

, and V. E. Thatcher. 1974. Monogenetic trematodes (Monopisthocotylea: Dactylogyridae) 

from freshwater fishes of Colombia, South America.—Journal of Helminthology 48:59-66. 

, and 1976. New monogenetic trematodes from freshwater fishes of western Colombia 

with the proposal of Anacanthoroides gen. n. (Dactylogyridae).—Proceedings of the Helmin- 

thological Society of Washington 43: 129-134. 

, and 1977. Phanerothecium gen. nov. and Fundulotrema gen. nov. two new genera 

of viviparous Monogenoidea (Gyrodactylidae), with a description of P. caballeroi sp. nov. and 

a.key to the subfamilies and genera of the family.—Instituto de Biologia, Publicaciones Es- 

peciales 4:53-60. 

—, , and R. J. Kayton. 1979. Neotropical Monogenoidea. 2. The Anacanthorinae Price, 


596 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


1967, with the proposal of four new species of Anacanthorus Mizelle & Price, 1965, from 
Amazonian fishes.—Acta Amazonica 9:355—361. 

————. , and 1980. Neotropical Monogenoidea. 3. Five new species from South 
America with the proposal of Tereancistrum gen. n. and Trinibaculum gen. n. (Dactylogyridae: 
Ancyrocephalinae).—Acta Amazonica 10:411-417. 

Mayes, M. A., D. R. Brooks, and T. B. Thorson. 1981. Potamotrygonocotyle tsalickisi, new genus 
and species (Monogenea: Monocotylidae) and Paraheteronchocotyle amazonensis, new genus 
and species (Monogenea: Hexabothnidae) from Potamotrygon circularis Garman (Chondrich- 
thyes: Potamotrygonidae) in Northwestern Brazil.—Proceedings of the Biological Society of 
Washington 94:1205-1210. 

Mizelle, J. D. 1936. New species of trematodes from the gills of Illinois fishes.—American Midland 

Naturalist 17:785—806. 

, and A. R. Klucka. 1953. Studies on monogenetic trematodes. XIV. Dactylogyridae from 

Wisconsin fishes.—American Midland Naturalist 49:720-733. 

, and D. C. Knitsky. 1967. Unilatus gen. n., a unique Neotropical genus of Monogenea.— 

Journal of Parasitology 53:1113-1114. 

, and . 1969a. Studies on monogenetic trematodes. XXXIX. Exotic species of Mono- 

pisthocotylea with the proposal of Archidiplectanum gen. n. and Longihaptor gen. n.—Amer- 

ican Midland Naturalist 81:370-386. 

, and . 1969b. Studies on monogenetic trematodes. XL. New species from marine and 

freshwater fishes.—American Midland Naturalist 82:417-428. 

, and J. W. Crane. 1968. Studies on monogenetic trematodes. XXXVIII. Ancyro- 

cephalinae from South America with the proposal of Jainus gen. n.—American Midland Nat- 

uralist 80:186—198. 

, and C. E. Price. 1964. Studies on monogenetic trematodes. XXVII. Dactylogyrid species 

with the proposal of Urocleidoides gen. n.—Journal of Parasitology 50:579-584. 

, and 1965. Studies on monogenetic trematodes. XXVIII. Gill parasites of the piranha 

with proposal of Anacanthorus gen. n.—Journal of Parasitology 51:30—36. 

Molnar, K., G. Hanek, and C. H. Fernando. 1974. Ancyrocephalids (Monogenea) from freshwater 
fishes of Trinidad.—Journal of Parasitology 60:914—920. 

Paperna, I. 1969. Monogenetic trematodes of the fish of the Volta basin and South Ghana.—Bulletin 
de 1’1.F.A.N. 31:840-880. 

Price, C. E. 1966. Urocleidus cavanaughi, a new monogenetic trematode from the gills of the 
keyhole cichlid, Aequidens maroni (Steindachner).—Bulletin of the Georgia Academy of Sci- 
ence 24:117—120. 

——. 1968. Diaccessorius, a new genus of Monogenea from the gills of an Amazon River te- 

leost.—Acta Biologica Venezuelica 6:84—89. 

, and W. A. Bussing. 1967. Monogenean parasites of Costa Rican fishes. Part 1. Descriptions 

of two new species of Cleidodiscus Mueller, 1934.—Revista di Parassitologia 28:8 1-86. 

, and 1968. Monogenean parasites of Costa Rican fishes. II. Proposal of Palombi- 

trema heteroancistrium n. gen., n. sp.—Proceedings of the Helminthological Society of Wash- 

ington 35:54—S7. 

, and T. E. McMahon. 1966. Monocleithrium, a new genus of Monogenea from an Amazon 

River teleost.—Revista di Parassitologia 27:22 1-226. 

,and W. J. Nowlin. 1967. Proposal of Dawestrema cycloancistrium n. gen. n. sp. (Trematoda: 

Monogenea) from an Amazon River host.—Revista di Parassitologia 28: 1-9. 

, and N. G. Romero. 1969. First account of a monogenetic trematode from Paraguay: Am- 

Pphocleithrium paraguayensis n. gen. n. sp.—Zoologische Jahrbuecher 96:449_452. 

, and E. A. Schlueter. 1967. Two new monogenetic trematodes from South America.— 

Journal of the Tennessee Academy of Science 42:23-25. 

Price, E. W. 1938. The monogenetic trematodes of Latin America.—Livro Jubilar Prof. Travassos, 
Rio de Janeiro, Brasil 3:407-413. 

Szidat, L. 1973. Morphologie and Verhalten von Paragyrodactylus superbus n. g., n. sp., Erreger 
eines Fischsterben in Argentinien.—Angewandte Parasitologie 14:1—10. 

Thatcher, V. E., and D. C. Kritsky. 1983. Neotropical Monogenoidea. 4. Linguadactyloides brink- 
manni gen. et sp. n. (Dactylogyridae; Linguadactyloidinae subfam. n.) with observations on 


VOLUME 96, NUMBER 3 597 


its pathology in a Brazilian freshwater fish, Colossoma macropomum (Cuvier).—Proceedings 
of the Helminthological Society of Washington 50:305-—311. 


(DCK) Department of Allied Health Professions and Idaho Museum of Natural 
History, Idaho State University, Box 8002, Pocatello, Idaho 83209; (VET) Insti- 
tuto Nacional de Pesquisas da Amazonia, Manaus, Brazil. 


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INFORMATION FOR CONTRIBUTORS 


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CONTENTS 


Nectocarmen antonioi, a new Prayinae, Calycophorae, Siphonophora from California 

Angeles Alvarifio 

A new Strombina species (Gastropoda: Prosobranchia) from the tropical western Atlantic 
~ Richard S. Houbrick 
Carenzia, a new genus of Seguenziacea (Gastropoda: Prosobranchia) with the description of a 
new species James F. Quinn, Jr. 
The lycoteuthid genus Oregoniateuthis Voss, 1956, a synonym of Lycoteuthis Pfeffer, 1900 
(Cephalopoda: Teuthoidea) Ronald B. Toll 
Parandalia bennei (Pilargidae) and Spiophanes lowai (Spionidae), new species of polychaetous 
annelids from Nazatlan Bay, Pacific coast of Mexico Vivianne Solis-Weiss 
Amphisamythia galapagensis, a new species of ampharetid polychaete from the vicinity of abyssal 

hydrothermal vents in the Galapagos Rift, and the role of this species in rift ecosystems 
Robert Zottoli 
A new scale worm (Polychaeta: Polynoidae) from the hydrothermal rift-area off western Mexico 
at 21°N Marian H. Pettibone 
Minusculisquama hughesi, a new genus and species of scale worm (Polychaeta: Polynoidae) from 
eastern Canada Marian H. Pettibone 

Arctodrilus wulikensis, new genus, new species (Oligochaeta: Tubificidae) from Alaska 

Ralph O. Brinkhurst and R. Deedee Kathman 
A contribution to the taxonomy of the Enchytraeidae (Oligochaeta). Review of Stephensoniella, — 


with new species records Kathryn A. Coates 
Distocambarus (Decapoda: Cambaridae) elevated to generic rank, with an account of D. crockeri, 
new species, from South Carolina Horton H. Hobbs, Jr., and Paul H. Carlson 
Distocambarus (Fitzcambarus) carlsoni, a new subgenus and species of crayfish (Decapoda: 
Cambaridae) from South Carolina Horton H. Hobbs, Jr. 
Disposition of three species of Oniscoidea from western Atlantic seashores (Crustacea: Isopoda: 
Holaphilosciidae and Philosciidae) George A. Schultz 
An unusual species complex in the genus Eurycope (Crustacea: Isopoda: Asellota) from the deep 
North Atlantic Ocean George D. F. Wilson 


Sponge-inhabiting barnacles (Cirripedia: Archaeobalanidae) of the Carolinian Province, south- 
eastern United States, with the description of a new species of Membranobalanus Pilsbry _ 
Victor A. Zullo and Jon D. Standing 

Zeugophilomedes, a new genus of myodocopine ostracode (Philomedinae) ah 
Louis S. Kornicker 4 
A new species of Munidopsis from submarine thermal vents of the East Pacific Rise at 21°N 


(Anomura: Galatheidae) Austin B. Williams and Cindy Lee Van Dover 4 1 


The Nearctic species of the Bezzia bivittata group (Diptera: Ceratopogonidae) ‘ 
Willis W. Wirth and William L. Grogan, Jr. 
A comparative study of selected skeletal structures in the seastars Asterias forbesi (Desor), A. 
vulgaris Verrill, and A. rubens L., with a discussion of possible relationships 


E. K. Worley and David R. Franz 5 


Notes on the frog genus Cycloramphus (Amphibia: Leptodactylidae), with descriptions of two 
new species W. Ronald Heyer 5 
Leptodactylus riveroi, a new frog species from Amazonia, South America (Anura: Lepto- — 
dactylidae) W. Ronald Heyer and William F. Pyburn © 
An annotated checklist of the fossil tortoises of China and Mongolia Charles R. Crumly 
Neotropical Monogenea. 5. Five new species from the Aruana, Osteoglossum bicirrosum Vandelli, 
a freshwater teleost from Brazil, with the proposal of Gonoclethrum n. gen. (Dactylogyridae: _ 
Ancyrocephalinae) Delane C. Kritsky and Vernon E. Thatcher 


(ISSN 0006-324X) 


Proceedings 
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Volume 96 29 December 1983 Number 4 


THE BIOLOGICAL SOCIETY OF WASHINGTON 


1983-1984 
Officers 
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Elected Council 
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PROCEEDINGS 
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PROC. BIOL. SOC. WASH. 
96(4), 1983, pp. 599-604 


THREE NEW SPECIES OF BRUEELIA 
(MALLOPHAGA: PHILOPTERIDAE) FROM 
THE MIMIDAE (AVES: PASSERIFORMES) 


N. Sandra Williams 


Abstract.— Three new species of Brueelia from the Mimidae are described: one 
from Oreoscoptes montanus, one from Toxostoma dorsale dorsale and one from 
Mimus polyglottos polyglottos. A key is provided for their separation. 


Of the 130 species of Brueelia recognized by Hopkins and Clay (1952, 1953, 
1955), none have been described from the Mimidae. The present paper concerns 
the Brueelia from this host family and is a step toward a comprehensive review 
of the entire genus. Thus far, Ansari (1956, 1957) and Williams (1981, 1982) have 
revised the species found on the Corvidae and Meropidae, respectively. For a 
synonymy of the Brueelia, see Williams (1981). 

This study deals with Mallophaga collected from 12 of the 35 recognized species 
of Mimidae. Gruson (1976) and A.O.U. (1957) were used for the nomenclature 
of the hosts. 

In the following ““Dimensions”’ sections, the numbers following each character 
represent its range, sample size, mean and standard deviation in mm. The “ab- 
dominal width” is the measurement of the widest abdominal segment which is 
the fourth, except for male B. polyglotta, where it is the fifth. 

Abbreviations for the location of the collections are BM(NH) (British Museum 
(Natural History)), UU (R. E. Elbel Collection, University of Utah), NMNH 
(National Museum of Natural History), and EEM (K. C. Emerson Entomology 
Museum, Oklahoma State University). 

The Brueelia from the Mimidae share the following characteristics with those 
from other hosts: ventral marginal carina interrupted medially, filiform antennae, 
prothorax small with lateral margins slightly concave, pterothorax broader than 
long with sides diverging and posterior margin evenly rounded, with tergal plates 
III and IV of the female separated medially. 


Key to the Brueelia from the Mimidae 


1. Setae on posterior edge of pterothorax (lateral to median) usually seven 


in number (four short, three long) in S-L-S-L-S-L-S pattern ...... montana 
— Setae on posterior edge of pterothorax (lateral to median) usually seven 
in number (one short and six long) in a S-L-L-L-L-L-L pattern ........ y) 


2. Male genitalia narrow with two sensilla on each side of mesosome and 
distal edges of mesosome with ridges. Parameres elongate. Margin of vul- 
val plate usually with eight setae of equal diameter on each side, three 
ALOVEranGeinNVeRDelOwWmre ets shea tre ce eet CULE ee le a ae dorsale 
— Male genitalia broad with two sensilla on each side of mesosome and distal 
edges of mesosome smooth. Parameres blunt. Margin of vulval plate usu- 


600 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


ally with 13 setae on each side, nine of wide diameter above four slender 
OMES: ace Rye 5 CON: A a Ee eet ee ee ee Ee nee ae polyglotta 


Brueelia montana, new species 
Figs. 4-6 


Type host.— Oreoscoptes montanus (Townsend). 

Brueelia montana is distinguished from other Brueelia species of the Mimidae 
in having four short and three long setae on the posterior margin of the pterothorax 
(lateral to median) in a S-L-S-L-S-L-S pattern. The male genitalia closely resemble 
that of B. dorsalis n. sp. but differs in being more narrow in the portion of the 
basal plate anterior to the parameres. 

Diagnosis.—Thimble-shaped head; ventral marginal carina interrupted medi- 
ally; dorsal marginal carina complete but indented medially; no dorsal anterior 
plate; antennae filiform and similar in both sexes; prothorax small with lateral 
margins slightly concave; pterothorax broader than long with sides diverging and 
posterior margin evenly rounded; abdomen elongate-oval, with tergal plates I- 
VII of male and female separated medially; male genitalia with slender basal plate; 
parameres narrow and short with proximal head simple; mesosome shield-like 
with two sensilla and ridges on distal edge; female vulval plate usually with seven 
setae of equal diameter on each side, three above and four below. 

Dimensions.— Male: Total length, 1.39-1.71 (5:1.55 + 0.1468); head length, 
0.32—0.39 (5:0.36 + 0.0324); head width, 0.25—0.34 (5:0.30 + 0.0418); prothorax 
width, 0.16—0.20 (5:0.18 + 0.0200); pterothorax width, 0.24—0.35 (5:0.30 + 
0.0482); abdominal width, 0.32-0.45 (5:0.39 + 0.0638). 

Female: Total length, 1.66—2.05 (18:1.87 + 0.1070); head length, 0.34—-0.52 
(18:0.40 + 0.0377); head width, 0.28-0.37 (18:0.33 + 0.0320); prothorax width, 
0.17-0.22 (18:0.20 + 0.0196); pterothorax width, 0.27-0.34 (18:0.32 + 0.0216); 
abdominal width, 0.36-0.51 (18:0.45 + 0.0428). 

Material Examined.— Holotype ¢ and allotype 2 from Oreoscoptes montanus, 
Ditto Dunes, Tooele Co., Utah, May 1969 (coll. E. and E. Branch, Dugway Proving 
Grounds), UU collection No. EE09054, deposited in NMNH,; paratypes 2: 66 7 
22, with same data; 2 °°, with same data except No. EE09107; 1 6 4 99, O. 
montanus, Vernon, Tooele Co., Utah, UU; 2 66 5 92, O. montanus, N. Skull 
Valley, Tooele Co., Utah, UU. 


Brueelia dorsale, new species 
Figs. 1-3 


Type host.—Toxostoma dorsale dorsale Henry. 

This species is distinguished from B. montana to which it is otherwise similar 
by pterothorax setation, by features of the male genitalia and female vulval plate 
and the shape of the head. The posterior edge of the pterothorax with one short 
and six long setae on each side. The male genitalia of B. dorsalis are narrow and 
have mesosomes with ridges on the distal edge. The margin of the vulval plate 
of females usually has eight setae of equal diameter on each side. The heads of 
both sexes are broadly triangular with a flattened anterior edge. 

Dimensions. — Male: Total length, 1.50-1.78 (38:1.64 + 0.0619); head length, 
0.36-0.42 (38:0.38 + 0.0113); head width, 0.35—0.39 (38:0.37 + 0.0094); pro- 


VOLUME 96, NUMBER 4 601 


CULIG 


Figs. 1-6. 1-3. Brueelia dorsale: dorsal and ventral views of (1) female and (2) male, and (3) male 
genitalia. 4-6. Brueelia montana: dorsal and ventral views of (1) female and (2) male, and (3) male 
genitalia. 


602 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


thorax width, 0.19-0.22 (38:0.20 + 0.0086); pterothorax width, 0.31—0.37 (38: 
0.33 + 0.0126); abdominal width, 0.48-0.55 (38:0.52 + 0.0238). 

Female: Total length, 1.47—2.10 (63:1.91 + 0.1318); head length, 0.38—-0.43 
(63:0.41 + 0.0094); head width, 0.37-0.45 (63:0.40 + 0.0117); prothorax width, 
0.20—0.26 (63:0.23 + 0.0095); pterothorax width, 0.32-0.41 (63:0.37 + 0.0142); 
abdominal width, 0.48-0.64 (63:0.58 + 0.0291). 

Material Examined.— Holotype ¢ (second specimen from the left as seen with 
compound microscope) and allotype @ (first specimen from the right with com- 
pound microscope) from Toxostoma dorsale dorsale, California, Mar 1939, (coll. 
Meinertzhagen), BM(NH) Collection, No. 13081 (slide #1 of 4), deposited in 
BM(NH); paratypes 27 66 57 °°, with same data; 1 6 7 °°, T. dorsale, Las Vacas, 
Coahuila, Mexico, UU; Other Material: from 7. rufum, 1 °, Urbana, Illinois, 
NMNH; | 2, John’s Island, South Carolina, NMNH; | 2, Gulfport, Mississippi, 
NMNH; from 7. curvirostre, 1 °, Chihuahua, Mexico, NMNH; 1 9, Las Vacas, 
Coahuila, Mexico, UU; from T. redivivum, 9 6é 13 9°, Pasadena, California, 
NMNH. In addition, K. C. Emerson has identified 1 ¢ 1 2 from 7. rufum, Co- 
lumbus, Ohio, and 1 2 from 7. rufum, Orient, New York, EEM. 


Brueelia polyglotta, new species 
Figs. 7-9 


Type host.—Mimus polyglottos polyglottos (Linnaeus). 

Brueelia polyglotta is distinguished from B. dorsale by a broad male genital 
plate, male genitalia with distal edges of the mesosome without ridges and blunt 
parameres, and the margin of the female vulval plate with 13 setae of varying 
diameters on each side. 

Dimensions.— Male: Total length, 1.27-1.58 (70:1.39 + 0.0070); head length, 
0.39-0.45 (70:0.41 + 0.0002); head width, 0.38-0.44 (70:0.40 + 0.0002); pro- 
thorax width, 0.20-—0.24 (70:0.23 + 0.0001); pterothorax width, 0.32-0.39 (70: 
0.35 + 0.0002); abdominal width, 0.42—0.58 (70:0.50 + 0.0007). 

Female: Total length, 1.46-1.98 (160:1.64 + 0.0102); head length, 0.41-0.48 
(160:0.44 + 0.0003); head width, 0.39-0.48 (160:0.43 + 0.0003); prothorax width, 
0.22-0.27 (160:0.24 + 0.0001); pterothorax width, 0.33-0.42 (160:0.37 + 0.0003); 
abdominal width, 0.45—0.62 (160:0.52 + 0.0012). 

Material Examined.— Holotype 4, and allotype ¢@ (first specimen on the right 
and middle specimen, respectively, as seen under the compound microscope) and 
paratype 2 (first specimen from the left as seen under compound microscope) 
from Mimus polyglottos polyglottos, Fairhope, Alabama, Sep 1930, (coll. Ms. W. 
M. Edwards), No. 18364 (slide No. | ofa set of 2), deposited in UMNH,; paratypes: 
13 66 15 92 with same data except Nos. as follows: 18354, 18355 (2 slides), 18357 
(2 slides), 18359, 18361, 18363, 18364 (2nd slide), 18366 (3 slides); from M. 
polyglottos, 2 66, Raleigh, North Carolina, UMNH; 6 22, Sonoro, Texas, UMNH; 
9 66 11 99, Alachua Co., Florida, UMNH; 5 4é 1 2, Savannah, Georgia, UMNH; 
1 2, Nashville, Georgia, UMNH; 1 6 3 9°, James Is., South Carolina, UMNH; 2 
66, Bloomington, Indiana, UMNH; 2 22, Dimmit Co., Texas, UMNH; 1 8, Ja- 
maica, West Indies; 1 6 1 2, Caja de Meurtos, Puerto Rico, UMNH; 2 44 2 92, 
Vieques Island, W.I., UMNH,; 6 46 14 99, State College, Mississippi, EEM; 2 6¢ 
2 °°, Bloomington, Indiana, EEM; 1 6, Cromwell, Oklahoma, EEM; | °, New 
Jersey, EEM; Other Material: from M. gilvus, 3 66 2 22, near Lethem, Rupununi, 


VOLUME 96, NUMBER 4 603 


Figs. 7-9. Brueelia polyglotta: dorsal and ventral views of (1) female and (2) male, and (3) male 
genitalia. 


British Guiana, BM(NH); 1 2, Aguachica, Magdalena, Columbia, BM(NH); from 
M. gundlachii, 15 66 46 22, Mariguana Is., Bahama Is., UMNH; 19 66 57 92, Great 
Inagua Is., British W.I., UMNH; | 636 3 9°, Ragged Is., B.W.I., UMNH; 1 8, 
Guantanamo, Cuba, UMNH; from Dumetella carolinensis, 2 22, Gamboa, Pan- 
ama Canal Zone, Panama, UU; 3 36 3 99, Ft. Collins, Colorado, EEM; 2 34 2 99, 
Orient, New York, EEM; 1 6 1 2°, Mongohela Co., West Virginia, EEM; from 
Melanotis hypoleucus, 1 6 1 2, Huehuetenango, Guatemala, UU; from Rampho- 
cinclus brachyurus, 2 °°, St. Lucia, W.I., BM(NH). 

Also, from Mimus longicaudatus, 6 66 5 °°, from Ecuador were identified but 
not included in the measurements because they were collected from a captive bird 
for which insufficient collection data were available. 


Acknowledgments 


I wish to thank Mr. Christopher Lyal of the British Museum (Natural History), 
Dr. Don R. Davis of the National Museum of Natural History, Dr. Robert E. 


604 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Elbel, Salt Lake City, Utah, and Dr. K. C. Emerson, Sanibel, Florida, for loan of 
material. Iam especially grateful to Dr. Emerson and Dr. Elbel for their assistance 
in the preparation of this manuscript. 


Literature Cited 


Ansari, M. A. E. 1956. A revision of the Bruélia (Mallophaga) species infesting the Corvidae (Part 
I).—Bulletin of the British Museum (Natural History) Entomology 4:369—406. 

—. 1957. A revision of the Bruélia (Mallophaga) species infesting the Corvidae (Part II).— 
Bulletin of the British Museum (Natural History) Entomology 5:143-182. 

American Ornithologists’ Union. 1957. Check-List of North American birds. Baltimore. Port City 
Press. 691 pp. 

Hopkins, G. H. E., and T. Clay. 1952. A check list of the genera and species of Mallophaga. British 

Museum (Natural History). London. 362 pp. 

, and 1953. Additions and corrections to the check list of Mallophaga.— Annals and 

Magazine of Natural History (12):6:434—-448. 

, and 1955. Additions and corrections to the check list of Mallophaga (II).— Annals 

and Magazine of Natural History (12)8:177-190. 

Gruson, E. S. 1976. Checklist of the world’s birds. New York. Quadrangle. 212 pp. 

Williams, N. S. 1981. The Brueelia (Mallophaga: Philopteridae) of the Meropidae (Aves:Coraci- 

iformes).— Journal of the Kansas Entomological Society 54:510-518. 

. 1982. The status of Brueelia superciliosa (Mallophaga: Philopteridae).— Journal of the Kansas 

Entomological Society 55:334. 


Biology Department, Simmons College, Boston, Massachusetts 02115. 


PROC. BIOL. SOC. WASH. 
96(4), 1983, pp. 605-622 


DESCRIPTION AND PHYLOGENY OF 
ISAACSICALANUS PAUCISETUS, N. GEN., N. SP., 
(COPEPODA: CALANOIDA: SPINOCALANIDAEB) 
FROM AN EAST PACIFIC HYDROTHERMAL 
VENT SITE (21°N) 


A. Fleminger 


Abstract.—A new genus and species of Spinocalanidae is described under the 
name Isaacsicalanus paucisetus. It was collected from the submersible R/V Alvin 
adjacent to the seafloor of the East Pacific Rise off the mouth of the Gulf of 
California. The copepod was observed swimming in a small swarm, apparently 
unispecific and all adult females, among concentrations of macrobenthic inver- 
tebrates typical of known hydrothermal vent communities. The new copepod’s 
most distinctive morphological features relative to other spinocalanids are the 
longer urosome, shorter antennule, and fewer setae on oral appendages. Phylo- 
genetic analysis supports the hypotheses that the new genus occupies a highly 
derived position within the Spinocalanidae and that the family is the most prim- 
itive within the superfamily Clausocalanoidea. 


Taxonomic reports on the recently discovered bathyal suspension-feeding mac- 
robenthos assemblages immediately adjacent to hydrothermal vents along oceanic 
spreading centers (Lonsdale 1977) reveal that virtually all the constituent species 
and their respective genera are new to science. The taxonomically unique qualities 
of these assemblages suggest that these species have persisted within a hydro- 
thermal vent-dominated habitat for appreciable periods of evolutionary time. 
Micro-invertebrates have also been represented; Humes and Dojiri (1980) de- 
scribed a new genus and species of thigmotactic siphonostome copepod found on 
the tentacular crown of tubicolous vestimentiferan worms from vent localities on 
the Galapagos rift. The present report considers a free-living planktobenthic cala- 
noid copeod that appears to be a member of a hydrothermal vent community on 
the East Pacific Rise at the mouth of the Gulf of California. 

The new form is morphologically most like Teneriforma Grice and Hulsemann, 
1967; several unique character states in the new form require minor emendation 
of Damkaer’s (1975) description of Spinocalanidae. 


Materials and Methods 


Thirty-two adult female individuals comprise the specimen series available for 
this study. They were collected adjacent to the seafloor at the East Pacific Rise 
several hundred kilometers west of Cabo Corrientes, Mexico; sampling details are 
presented below as part of the species’ description. Initial preservation was by 
freezing in sea water at — 20°C following transfer to the surface. The specimens 
were stored frozen at —70°C for about 10 weeks in the laboratory before transfer 
to formalin. They were immersed in glycerol for study. Internal organs are mostly 


606 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


indistinguishable, presumably a consequence of freezing at relatively high tem- 
peratures and precluding observations on the nature and contents of the midgut 
as well as limiting evaluation of ovarian development. 

All measurements were made of specimens immersed in glycercol. Figures were 
drawn with the aid of a camera lucida. Habitus views were drawn from specimens 
in which soft tissues were removed by KOH digestion and the cuticle lightly 
stained with chlorozol black. 

List of abbreviations used in text and figures.—A1, antennule; A2, antenna; B1, 


coxal segment of appendage; B2, basal segment of appendage; Lil, 2... , medial 
lobe(s) on protopod of cephalic appendage; Md, mandible; Mx1, maxillule; Mx2, 
maxilla; Mxp, maxilliped; Pl, 2, 3 ..., paired swimming legs 1, 2, 3 ...; Rel, 
2...,exopod, segment 1,2...; Ril, 2..., endopod, segment 1, 2...; Sel, 2 


..., lateral spine on exopod of swimming leg; Si, seta; St, terminal spine on 
distalmost exopodal segment of swimming leg; ThI, II, III, lV, V pedigers (=tho- 
racic segments, Bowman 1976:187) bearing swimming legs 1, 2,3. . . respectively. 


Family Spinocalanidae Vervoort, 1951 


Diagnosis (emended from Damkaer 1975).—Calanoids with P1 Re trimerous. 
Pl Rel with O or | Se, Re2 with O or 1 Se, Re3 with 1 Se, 1 St and 4 Si; Ri with 
one segment and 3 to 5 Si. Re trimerous in P2—4, Re3 with 3 Se, 5 Si, and one 
finely serrate St; lateral margin of Re3 not serrate. P2 with bimerous Ri, Ri2 with 
2 medial, 2 terminal and O or | lateral setae. P3—4 with trimerous Ri, Ri2 with 
1 medial seta, Ri3 with 2 medial, 2 terminal and O to 2 lateral setae. In female, 
cephalosome and ThI separate, ThIV and V usually separate, ThV distal end 
sometimes prolonged, apex may be pointed; P5 absent; Al with segments 8 and 
9 fused, 24 and 25 articulating or fused; A2 Re usually equal to or longer than 
Ri; setation of mouthparts and other details given in Tables | and 2. 


Tsaacsicalanus, new genus 


Diagnosis.—Robust spinocalanid; female with elongate urosome about half 
length of prosome. Short Al about as long as prosome. Rostrum short, lobate 
without any semblance of bifurcation. Mx1 with B2 fused to proximal part of Ri 
and bearing 4 Si; Li 2 with 1 seta, Li3 lacking Si. Distal Si of Mx2 and Mxp 
unarmed except for very fine setules on proximal side. Mxp B1 with 2 Si. Pl Rel 
and 2 lacking Se. P1 Ri with 5Si and prominent shoulder (lateral swelling produced 
anterodistad, knob-like) at segment’s midlength and bearing spinules concentrated 
at apex. P2—P4 with Re3 Se3 about twice as long as Sel and 2. A2 Rel without 
Si, Re2 and 3 not fused, Re2 with 1 Si. Posterior surfaces of P2—P4 without spines. 

Etymology.—The generic term [saacsicalanus is masculine and a combination 
of the late John D. Isaacs’ surname and -ca/anus, the suffix of many calanoid 
copepod genera. The name is proposed in memory of John D. Isaacs, a pioneering 
leader in oceanography, to acknowledge his many scientific contributions, the 
brilliant and the outrageous, and for his unwavering and most generous support 
of zooplankton research at Scripps Institution of Oceanography (SIO). It is also 
a personal expression of gratitude for the many kindnesses he showed me during 
20 years of collegial association at SIO. 

The type-species of the new genus is by original designation, J. paucisetus, 
described below. 


VOLUME 96, NUMBER 4 607 


A 


4 
Mu 


Fig. 1. Jsaacsicalanus paucisetus. Adult female. A, Habitus, dorsal; B, Habitus, lateral; C, Forehead 
and rostrum, ventral; D, Rostrum, anterior; E, Genital segment, ventral; F, Genital segment, lateral; 
G, Furca, dorsal. 


DEFG 


Key to Adult Females of Spinocalanid Genera 


1. Pl Rel with Se; P2 to P4 with one or two rows of spines on posterior 


SUITPACS WI ROWSE, UIE NOI viele VS 2 anne MPO SRI CT 0 SURE AVA ee, Pee Rye 2) 
— P1 Rel without Se; P2 to P4 lacking row of spines on posterior surface 
OL TRVD SPSS, Aine ee ee ene aD Oe ee ere OMS ny MON Se Ores eee ty a 3 


ee RA 8 ei Mved ante tamed yar een Eee ae Monacilla Sars, 1905 


INOStHUIMEAOSEM tain od. es ee ed BESS See Spinocalanus Giesbrecht, 1888 
3. Rostrum absent. Prosome at least 4 times length of urosome, caudal ramus 

length not more than 1.5 times width ......... Mimocalanus Farran, 1908 
— Rostrum present but not bifurcate. Prosome no more than 3 times length 

of urosome. Caudal ramus length about 2 times width ............... 4 
4. Al, extended, reaching to anal segment, A2 with endopod longer than 

exopod, Mxp BI with5 Si ...... Teneriforma Grice and Hulsemann, 1967 
— Al, extended, reaching to ThV, A2 with exopod longer than endopod, 

INVA cpap ss ava il 28 Sie yes cay gs hag ea ospesbelialhey aac. cheephyaatea he ola Be Tsaacsicalanus, gen. n. 


TIsaacsicalanus paucisetus, new species 
Bigs. lee S 


Measurements.— Adult female total length to distal end of caudal ramus, range 
2.44 to 2.56 mm, xX 2.501 + 0.0259 mm (95% C.L., N = 10, s = 0.0368). 

Prosome:urosome length ratio, measurements taken along mid-sagittal plane 
from apex of forehead to hinge between prosome and urosome and from hinge 
to distal end of right caudal ramus; range 2.03 to 2.19:1, median 2.15:1 (N = 10). 


608 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


ABCERG 


Fig. 2. Jsaacsicalanus paucisetus. Adult female. A, Right Al segments 19 to 25, dorsal; B, Right 
A2, posterior; C, Right Mnd, posterior; D, Right Mnd gnathobase, posterior; E, Right Mxl, posterior; 
F, Right Mx2, anterior; G, Left Mxp, anterior. 


Description and diagnosis of adult female.—WHabitus (Figs. 1A, B), forehead 
broadly rounded, prosome short, about 2.15 x length of urosome; in Teneriforma 
naso (Farran, 1936) prosome is at least 2.5 x length of urosome (measured as 
above using illustrations in Farran 1936, Grice and Hulsemann 1965 and Wheeler 
1970); ThV distal end produced and terminating in a sharp point; Al short, 
extended posteriorly not reaching beyond ThV. Caudal ramus about twice as long 
as wide (Fig. 1G) bearing 4 terminal setae and | short medial seta at about 
midlength of ramus. Cephalosome and ThI separated, ThIV and V incompletely 
separated, remnant of line of articulation appearing on dorsal side. Rostrum in 
lateral view short knob-like; in frontal and ventral views (Figs. 1C, D) a short 
simple lobate process. Genital segment with relatively large seminal receptacles 
curving anteriorly and swollen distally (Figs. 1E, F). 

Al with 23 articulating segments, segments 8 and 9 and 24 and 25 fused (Fig. 
2A). A2 (Fig. 2B) with Re about 1.3 times longer than Ri. Re with total of 9 Si; 


VOLUME 96, NUMBER 4 609 


O.2mm 
ANBED 


Fig. 3. Isaacsicalanus paucisetus. Adult female. A. Right P1, anterior; B, Right P2, posterior; C, 
Right P3, posterior; D, Right P4, posterior. 


. Rel lacking Si, Re2 with 1 Si, remnant of apical segment (Re8) with 3 Si, distinctly 
separate from preceding segment, Re7, by visible remnant of intersegmental su- 
ture; Re7 with | Si. Re in Teneriforma with total of 11 Si. 

Md palp (Fig. 2C) with setation similar to that in Teneriforma and Mimocala- 
nus. Left and right gnathobase similar, with 8 major teeth of roughly equal size, 
a small intercalary tooth between the second and third and the third and fourth 
major teeth and a slender, short Si (Fig. 2D). Ventralmost tooth slender, spiniform, 
set off from others by deeply cut angular notch; remaining teeth distally bifid; 3 
proximalmost teeth bordered by minute denticles. 

Mx! (Fig. 2E) with proximal portion of Ri totally fused to B2 and bearing 4 
Si; these parts also fused in Teneriforma but combination bearing 10 setae; Ri 
and B2 not fused elsewhere in family. Li2 with 1 seta, Li3 much smaller than Li2 
and lacking Si; elsewhere in family Lil and 2 bear a total of 6 or more Si. 

Mxp (Fig. 2G) with B1 bearing 2 Si, elsewhere in family this segment with 5 
or more Si. Si on distalmost segments with minute setules. Ri segments 2 and 3 
with 2 Si, Ri4 with 3 Si. 

P1 (Fig. 3A) with B2 Si strongly curved as in Monacilla; this Si straight in 
Teneriforma. Lateral shoulder of Ri located at about midlength of segment, strong- 
ly produced anterodistad in knob-like process with apex bearing numerous spi- 
nules; Ri bearing 5 Si as in Monacilla. Both Rel and Re2 lacking Se as in Te- 
neriforma described by Farran (1936); Grice and Hulsemann (1965) show P1 Re2 
in Teneriforma with | Se. 


610 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


P2 to 4 (Figs. 3 B, C, D). Re3 of P2 to P4 with Se3 about twice as long as Sel 
and Se2. Re2 of P2 to P4 with Se about 1.5 x longer than Rel Se. P2 Ri2 with 
1 lateral Si; P3 and P4 Ri3 with 2 lateral Si as in Teneriforma. P2 to P4 Ri2 
disto-lateral corner rounded as appears to be the condition in the Australian 
Teneriforma naso described by Farran (1936) but not in 7. naso described by 
Grice and Hulsemann (1965) where this corner is produced in a short pointed 
denticle as it is elsewhere in the family. 

Etymology.—The specific name, paucisetus, literally ‘few setae,’ signifies the 
reduced number of setae on oral appendages of the new species relative to other 
species of the family. 

Types.—Female holotype, TL 2.63 mm, USNM No. 195066; 15 paratypes, 
USNM No. 195067, all taken together at the type-locality. 

Sampling data and type locality.— All specimens obtained in one collection 
made at 20°49.05’N, 109°06.40'W on 23 April 1982 at R/V Alvin Dive no. 1217, 
Sample no. 1217-3B, during Oasis Expedition to the East Pacific Rise. The col- 
lection was made by Roberta Baldwin, SIO, with aid of a 4-chambered slurp gun 
respirometer (Smith and Baldwin, in press) within | m of the sea floor at a depth 
of 2600 m. The copepods were swimming in a tightly clustered swarm of several 
thousand individuals just above a small depression known as “Holger’s Hole” 
about | m in diameter and | m deep formed in collapsed pillow lava. With regard 
to the macrobenthos at the site, I quote from R. Baldwin’s field notes: ““One side 
of the depression is bounded by a dense population of Calyptogena magnifica 
Bossand Turner (Vesicomyidae) and a small patch of Riftia pachytila Jones (Ves- 
timentifera); the bottom, littered with broken lava pieces, is inhabited by the 
clams and Munidopsis sp. (Galatheidae). Water temperatures in the depression 
range from 5° to 15°C. The collapsed pillow lava site is located in an area known 
as “clam acres” (2613 m depth) where dense populations live in the mineral-rich 
waters of the hydrothermal vents off Baja California. The area is located 240 km 
south of the Baja peninsula and 21° north of the equator on the East Pacific Rise, 
an area of seafloor spreading where bottom-water temperatures range from 2.2°C 
over the clam fields to an excess of 350°C in the stalagmite chimneys known as 
‘black smokers.’’ The collection was incubated in situ in the respirometer for 
about 48 hours before being brought to the surface and frozen. Randomly selected 
subsamples from the copepod swarm, totalling 32 individuals are all adult females 
of the new species. 


Phylogenetic Relationships among Spinocalanid Genera 


The family Spinocalanidae was established to accommodate three genera, Spi- 
nocalanus, Monacilla and Mimocalanus, that Vervoort (1951) appropriately re- 
moved from the family Pseudocalanidae (=Clausocalanidae Giesbrecht in Bow- 
man and Abele 1982). Knowledge of spinocalanid diversity and distribution 
accumulated rapidly during the ensuing 20 years in response to widespread oceanic 
exploration and increased use of opening-closing nets sampling meso- and bath- 
ypelagic depths where spinocalanids tend to be most commonly encountered. 
Damkaer (1975) brought these advances into full focus by virtue of his thorough 
review of the Spinocalanidae. He recognizes 32 species and 4 genera: 19 species 
in Spinocalanus, 4 in Monacilla, 8 in Mimocalanus and 1 in Teneriforma. Damkaer 


VOLUME 96, NUMBER 4 611 


apparently regarded gaps in morphological knowledge of many spinocalanids 
sufficient reason to preclude consideration of phylogenetic relationships, but he 
suggested that Monacilla and Spinocalanus comprise a group and that Mimo- 
calanus and Teneriforma comprise another group within the family. 

Knowledge of species recognized by Damkaer (1975) has not advanced, but the 
new genus /saacsicalanus provides additional infrastructure to the family and, in 
apparently expanding the family’s range of habitats to a planktobenthic niche 
within the uniquely warm bathyal waters of a hydrothermal vent community, 
enhances interest in seeking its phylogenetic relationships. Classification of [saac- 
sicalanus, based solely on adult female morphology, derives from analysis of more 
than 30 characters. These characters reveal a number of apparent synapomorphies 
among spinocalanid genera. Following procedures discussed at length by Eldredge 
and Cracraft (1980:19-85) and briefly outlined by Sanders (1981:96-98), the 
available data set has been used to hypothesize a first approximation of phylo- 
genetic relationships among the spinocalanid genera. In this exploratory attempt 
to order the genera phylogenetically, character states representing other spino- 
calanid genera were collated from published accounts of the species. Two criteria 
were stressed in hypothesizing apomorphies and deciding upon the transformation 
sequence of character states (Table 1), namely, uniqueness and relative abundance 
of meristic features. For repetitive segments or their armature I have assumed 
that within a monophyletic lineage the highest numbers of setae and segments 
are plesiomorphic, regardless of frequency of occurrence, and that reductions are 
apomorphic. The most primitive state for each character, found in one or more 
members of a genus was selected to represent that genus. 

For some characters, determining the sequence of character state transforma- 
tions is readily obvious from the first principles of apomorphy by reduction. Many 
calanoid morphological features are generally conservative and the plesiomorphic 
condition widespread. Derived states are immediately recognizable when meristic 
features are reduced by developmental modifications, e.g., by fusion of adjacent 
segments, loss of the fifth pair of legs in the females, reduced number of setae on 
certain appendages, the absence of the rostrum or its failure to bifurcate. The 
direction of transformation in other characters may not be obvious. Solution 
requires outgroup comparison, 1.e., comparative analysis using as references closely 
related antecedent and derived taxa in which the character states in question are 
ummistakably primitive or derived relative to the states under consideration. 

Selection of an appropriate outgroup for the Spinocalanidae is not immediately 
obvious. As is typical of supraspecific calanoid taxa, the present concept of Spi- 
nocalanidae is a constellation of mostly plesiomorphic states that distinguishes it 
from the other 10 families comprising the superfamily Clausocalanoidea (Bowman 
and Abele 1982). Assuming that convergent reversals have not contributed to the 
formation of spinocalanids, their ancestry must derive from calanoids with char- 
acter states as follows: 


Bifurcate rostrum 

P2-—4 with Re3 bearing 3 Se and 5 Si 

Al with 25 articulating segments or with 8 and 9 fused 

Left and right members of male P5 biramous 

Trunk segmentation and genital segment morphology: 1.e., cephalosome and 


OV RwWN So 


612 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Table 1.—Catalogue of characters. 


Derived 


Char- character 
acter States recognized and state 
code Character proposed sequence of transformation* no. 
*Character state symbols: 0 = primitive condition, derived states coded by integers; 
+ + — hypothesized sequentially derived autapomorphies; 
— > hypothsized irregularly derived autapomorphies; 
| hypothesized autapomorphies reversing sequence. 
SA PI Ri 1-segmented with lateral shoulder, 3 to 5 Si 01 
SB PI Re3 with 4 Si 02 
SC P2-4 Re3 with 3 Se and 5 Si 03 
SD P5 lacking in female, endopod may be present in 
male but lacking setae 04 
SE AI segments 8 and 9 partly to totally fused 05 
SF cephalic appendages show reductions in size and setation in male 
relative to female 06 
A rostrum bifurcated—0; absent—07; uniramous—08 
hypothesized transformation 07-—0-08 07, 08 
B’ Al segments 24 and articulated—O; fused—09 09 
25 
C  A2 Ril setae no. 2—0; 1—10 10 
D_ A2 Ri? setae no. 18—O; 16—11; 15—12 11, 12 
0 
hod 
[ele 
E  A2 Rel setae no. 1—0; 0—13 13 
F  A2 Re? setae no. 4—0; 3—14; 1—15; 0—16 14, 15, 16 
0-14-15 
{ 
16 
G_ A2 Re7 setae no. I1—0; 0—17 17 
H Mand, no. teeth S8—0; 7—18 18 
I Mnd, B2 setae no. 4—0; 3—19 19 
J Mnd, Ri setae no. 14—0; 13—20; 1/—21 20, 21 
0-20-21 
K Mx! Re setae no. 12—0; 11—22; 10—23; 9—24 ph Pray De 
0-22-23 
| 
24 
L MxlRi+B2setaeno. 30—0; 21—25; 20—26; 13—27; 8—28 MS), 25, 27), 
28 
0-27-28 
{ 
25 
{ 
26 
M Mx! Ri and B2 articulated—0; fused—29 29 
segments 
N Mx! Li3 setae no. 4—0; 2—30; 0—31 30, 31 
0-30-31 
O Mx! Li2 setae no. 6—0; 5—32: 4—33; 1—34 32, 33, 34 
0-33-34 
{ 


32 


VOLUME 96, NUMBER 4 613 


Table 1.—Continued 


Derived 
Char- character 


acter States recognized and state 
code Character proposed sequence of transformation* no. 
P Mx! Lil setae no. 14—0; 13—35; 12—36; 10—37 35, 36, 37 
0-35-37 
{ 
36 
Q Mx2 Lil setae no. 6—9; 5—38; 3—39 38, 39 
0-38 
| 
39 
R  Mx2 Li2 setae no. 3—0; 2—40 40 
S  Mx2 Li5 setae no. 4—0; 3—43 43 
T Mx2 Ri setae no. 7—0; 5—41; 4—42 41, 42 
0-42 
{ 
41 
U  Mxp BI setae no. 10—0; 9—44- 5—45; 2—46 44, 45, 46 
0-44-4546 
V Mxp B2 setae no. 5—0; 4—47 47 
W Mxp Ril setae no. 5—0; 4—48; 3—49; 2—50 48, 49, 50 
0-48-49 
{ 
50 
X  Mxp Ri2 setae no. 4—0; 3—51; 2—52 51, 52 
0-51-52 
Y Mxp Ri3 setae no. 3—0; 2—53 53 
Z  Mxp Ri4 setae no. 4—0; 3—54 54 
AA P!1 B2Si shape curved—0O; straight—55 55 
BB PI! Rel spine no. 1—0; 0—56 56 
CC PI Re2 spine no. 1—0; 0—57 57 
DD PI! Ri setae no. 5—0; 4—58 58 
0 
| 
58 
EE P2-—4 spines on post. yes—0; no—59 59 
face of rami 
FF P3—4 Ri2 outer pointed—0O; rounded—60 60 
distal corner 
GG_ TRV posterior rounded—0; pointed—61 61 


comer 


ThI not fused, ThIV and ThV not fused; female abdomen with 4 segments 
and genital segment with large vertically oriented seminal receptacles 

6. Appreciable sexual dimorphism modifying appearance of male’s prosome, 
Al, and setation of cephalic appendages 


Spinocalanidae is the only family of the Clausocalanoidea with species exhib- 
iting states 2 and 4 above in combination (Monacilla spp. and Spinocalanus spp.). 
Additional plesiomorphic states within the family (e.g., numbers 3 and 5 above, 
oral appendages with highest number of typical setae within superfamily) mark 


it as containing the most primitive taxa within the superfamily. 


614 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Within the Clausocalanoidea aetideid genera Aetideopsis Sars, 1902, Pseudo- 
tharybis T. Scott, 1909, and, to a somewhat lesser extent, Bradyidius Giesbrecht, 
1897, came close to agreeing with the five character states characteristic of Spi- 
nocalanidae listed above. They are, however, apomorphic to spinocalanids in 
characters 2 and 5; 1.e., relative to spinocalanids they all show a loss of 1 Si on 
Re3 of P2—P4 and their thoracic segmentation includes partial to total fusion of 
cephalon and ThI as well as ThIV and ThV or pronounced foreshortening of 
ThV. Based on all characters analyzed in this study (Table 1) it is conceivable 
and even likely that these three aetideid genera derive from a Monacilla-like 
ancestor. The search for a closely related outgroup differing from spinocalanids 
only in a plesiomorphic direction was extended to other superfamilies. Looking 
beyond the Clausocalanoidea, the family Megacalanidae of the Megacalanoidea 
(aiso containing Calanidae, Calocalanidae, Paracalanidae, and Mecynoceridae; 
Bowman and Abele 1982) appears to be the most promising candidate in that the 
majority of characters I examined (Table 1) are symplesiomorphic or synapo- 
morphic to spinocalanids and the remainder are plesiomorphic. Two possible 
anomalous characters, |., a noncatalogued condition of asymmetry in adult males 
and, 2., character L, require special consideration, however, before designating 
the Megocalanidae as the outgroup of choice for determining the direction of 
transformation sequences (i.e., polarity of morphoclines) in spinocalanids. 

Among the males of Spinocalanus the orientation of asymmetries found in the 
Al and P5, and presumably the position of the genital pore, may appear on the 
left or the right side of the body depending upon the species (Damkaer 1975:14). 
To my knowledge asymmetries in megacalanid males do not vary between or 
within the species. Interspecific reversals in asymmetry, however, are known to 
occur in Clausocalanus (Frost and Fleminger, 1968), Calocalanus Giesbrecht, 
1888 (Andronov 1973) and the genital opening, typically on the left side in Cal- 
anoida, occurs on the right side of the urosome in Mecynocera Thompson, 1888 
(Andronov 1973). I have infrequently observed intraspecific reversals of asym- 
metry in the P5 of males of Eucalanus pileatus Giesbrecht 1888 inhabiting south- 
east Asian waters (unpublished). Variation in male asymmetries is a well known 
feature of the Metridiidae (superfamily Augaptiloidea, a likely source of the Me- 
gacalanoidea). Thus, reversal in the asymmetry of spinocalanid males clearly is 
of no special phylogenetic significance vis ad vis the Megacalanidae. Reversed 
asymmetry in males may be an old synapomorphy of the Augaptiloidea, Mega- 
calanoidea, Clausocalanoidea and Eucalanoidea, or in view of its sporadic oc- 
currences merely a number of convergent autapomorphies. 

The second exception to the evidence favoring Megacalanidae as the most 
suitable antecedent outgroup for Spinocalanidae is the number of setae on Mxl 
Ri plus B2 (Character L, Table 1). I have not found any species of calanoid to 
match the total of 30 setae on these segments occurring in Mimocalanus crassus 
(Park 1970:479, Fig. 5). Within Spinocalanidae setal count values range from 21 
or, less frequently, 20 setae in Spinocalanus (Park 1970: Figs. 27, 47, 73; Damkaer 
1975: Figs. 15, 48, 71, 122), 18 to 20 in Monacilla (Farran 1908: Pl. 1, Fig. 17; 
Scott 1909: Pl. II, Fig. 14; Sars 1925: Pl. XI, Fig. 9), 19 to 30 in Mimocalanus 
(Tanaka 1956: Fig. 13d; Vervoort 1957: Fig. 13c; Park 1970: Fig. 5), 13 in Te- 
neriforma (Grice and Hulsemann 1965: Fig. 8F), and 8 in Isaacsicalanus (Fig. 
2E). The MxI has not been characterized for a number of described spinocalanids. 


VOLUME 96, NUMBER 4 615 
Table 2.—Apparent synapomorphies among spinocalanid genera. 
Char- 
acter Mona Spino- Mimo- Teneri- Tsaacs!- Rever- 
code Character cilla calanus calanus forma calanus sals 
Females 
bifur- 
A Rostrum cated absent uniramous 
B Al segments 24 and 25 articulated fused 
C A2 Ril setae no. 1 py Dp; 2 2 x 
D A2 Ri2 setae no. 15 16 16 15 18 x 
E A2 Rel setae no. 1 1 0 1 0 x 
F A2 Re2 setae no. 4 3 0 3 1 x 
G A2 Re7 setae no. 1 1 1 0 1 x 
H Mnd no. teeth 8 8 8 7 8 x 
I Mnd B2 setae no. 4 4 3 3. 3 
J Mnd Ri setae no. 14 14 14 13 Ll 
K Mx! Re setae no. 12 11 11 9 10 x 
L Mx! Ri + B2 setae no. 20 21 30 13 Sine 2 x 
M Mx! Ri and B2 segments _.. articulated 0... fused 
N Mx! Li3 setae no. 4 4 4 2 Ome 
O Mx! Li2 setae no. 5 6 4 4 1 x 
P Mx! Lil setae no. 12 14 13 10 TO 3x 
Q Mx2 Li] setae no. 6 6 6 3 Dae nee ax 
R Mx2 Li2 setae no. 3 3 3 3 2 
S Mxz2 Li5 setae no. 4 4 3 3 3 
T Mxz2 Ri setae no. 5 7 4 4 4 x 
U Mxp BI setae no. 10 9 9 5 Dieta: 
V Mxp B2 setae no. 5 5) 5 4 4 
W Mxp Ril setae no. 5 4 4 2) 3 x 
x Mxp Ri? setae no. 4 4 4 3 Din ly 
Y Mxp Ri3 setae no. 3 3 3 2 I); eee 
Z Mxp Ri4 setae no. 4 4 4 4 3 
Ret ech ee curved uu... straight Xx 
AA Pl B2 seta curved 
BB P1 Rel spine no. 1 1 0 0 0) 
CC P1 Re2 spine no. 1 1 1 1 0 
DD P1 Ri setae no. 5 4 4 5 x 
EE P2—4, spines on post. VG Siete no ay, 
surfaces of rami 
FF R8=dRu2outer! PN) 0 wee ee Pointed est ea. aut rounded 
distal corner 
GG MEH SspoOstemOne i) 9) Lite ee ROUNGCC 2a ee eed pointed 
corner 
Males 
HH SINS Cath PLENARY biramous .......... uniramous ? qv 


Mx! Ri plus B2 may bear 21 setae in Clausocalanus (Frost and Fleminger 1968: 
Pl. 6, Fig. c) and in metridiids (Giesbrecht 1892: Pl. 32, Fig. 4). Calanids do not 
appear to exceed 19 setae, whereas megacalanids, also the outgroup of choice for 
calanids (in prep.), have 17 or fewer setae (Sars 1925: Pl. I, Fig. 11; Pl. III, Fig. 


616 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


: 
S 
: 8 
x Y 
s : 
R 8 
x 
6 
58158 60 
55 57 
5I 54 
50 52 
45 49 
39 46 
33 49 SS 
30 38 S 
27 Pa S 
24 Sip) eS 
20 28 
ig 23 59—4—59 
7 21 584—58 9 
14 4 15 56—+—56 & 
2 12 i313 43-4—43 3S 
42 42 = 
8 59—4—59 35 § 
S 56—}— 56 33 eS 
S 53 i9—+t—19 
S 47 16 25 
43—1— 43 13 13 l4a—t—14 
4 42—4— 42 
36 37 48 
—32 29 44 
26 19 Ig 22 
l2 2 09 I 
10 08 o7 
06 
05 
04 
03 
02 
Ol 


Fig. 4. Dendrogram to summarize distribution of derived character states among spinocalanid 
genera; derived states defined in Table 1 and discussed in text. 


7; Pl. IV, Fig. 7; Bjornberg 1968: Figs. 7, 27, 48, 57, 66). The low number of 
setae on these segments of Mxl in megacalanids may be an autapomorphy in the 
known bathypelagic species that appear to be adapted for predation, carrion feed- 
ing or large particulate dentrital feeding. Considering the incomplete state of 
knowledge on Mx] setation in the families in question, the 30 setae found on the 
Mx! Ri plus B2 in ™. crassus is tentatively regarded to be the plesiomorphic 
condition for Megacalanoidea and Clausocalanoidea. 

In summary, I have hypothesized a megacalanid-like ancestry for the Spino- 
calanidae and present the direction of character state transformations, 1.e., mor- 
phoclines, and proposed synapomorphies in Tables | and 2, respectively. 

As noted above, morphocline direction may be determined from the first prin- 


VOLUME 96, NUMBER 4 617 


Fig. 5. Cladogram of hypothesized synapomorphies in transformation sequences whose direction 
is proposed on the basis of first principles of calanoid copepod morphology. Further discussion in 
text. 


ciple of apomorphy by reduction in several instances. For example, the primitive 
condition of articulating segments of appendages or trunk are replaced autapo- 
morphically by fused combinations of adjacent segments as in the case of Nos. 
A and M (Table 2). Another assumed plesiomorphy is the bifurcate rostrum 
produced in a pair of slender processes as found in Monacilla. Complete loss of 
the rostrum (Character No. A) is synautapomorphic in Spinocalanus and Mim- 
ocalanus. Extension of the base of the rostrum into a single nonbifurcate process 
constitutes a second synautapomorphy in Jeneriforma and Isaacsicalanus. 


Results of Phylogenetic Analysis 


In the process of classifying Jsaacsicalanus 33 intrafamilial and six interfamilial 
character states were determined by microscopic examination of available adult 
females. States in the other spinocalanid genera were collated from published 
accounts of the species (Damkaer 1975; Farran 1908, 1936; Giesbrecht 1892; 
1971: Grice and Hulsemann 1965, 1967; Park 1970; Scott 1909; Sars 1903, 1925; 
Vervoort 1951, 1957; Farran and Vervoort 1951; Tanaka 1956; Wheeler 1970). 
Transformation sequences were proposed, generally on the basis of the first prin- 
ciple of calanoid copepod morphology and confirmed by outgroup comparison 
with Megacalanidae as described by Sars (1925), Sewell (1947), Bjornberg (1968), 
and my unpublished observations on Megacalanus and Bathycalanus. Character 
states and proposed transformation sequences are presented in Tables | and 2. 
States designated primitive indicate primitiveness of that state in that taxonomic 
category. This simplifying assumption is necessary due to the general lack of 
character surveys on calanoid families and genera as well as the logistic difficulties 
of obtaining specimens for study of seemingly critical species. In Table 1 all 
characters have been coded by letters, all derived, i1.e., autapomorphic, states are 
coded by integers. 

All derived states were used to construct a dendrogram (Fig. 4), following 
Johnson (1982) that depicts the distribution of derived states among genera of 


618 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


5 v? 6 
‘i \ alt 
All | a” ol ce ie° 


0) 0 0) G 
ea Cs 


ZwoxAscD 


™ @ 


es Se al aa Os 


Fig. 6. Cladogram of hypothesized synapomorphies where criterion in meristic states is identical 
numbers. Transformation sequences determined by outgroup comparison; further discussion in text. 


the Spinocalanidae. States 01 to 06 are familial and set Spinocalanidae apart from 
other families of the Clausocalanoidea. The remaining states provide compelling 
evidence of the distinctiveness of the five recognized genera and of the apparent 
synapomorphies that link Spinocalanus to Mimocalanus and Teneriforma to 
Isaacsicalanus. The handful of states appearing in more than one column, marked 
by the horizontal bar and integers appearing on both sides of the column, provide 
no indication of phylogenetic relationsips suggested by the analyses of proposed 
synapomorphies presented below. 

Potential synapomorphies are presented in Table 2. Initially troubled by the 
seemingly considerable taxonomic distance between the outgroup Megacalanidae, 
a primitive member of the Megacalanoidea, and Spinocalanidae, a primitive 
member of the Clausocalanoidea, I utilized sets of synapomorphies chosen by 
three different criteria to construct three separate cladograms for comparison. 

The first cladogram uses only those proposed synapomorphies (Table 2) whose 
transformation sequences appear reasonable and unequivocal solely on the basis 
of first principles of calanoid copepod morphology, i.e., the readily apparent 
transformation sequences B, M, BB, HH, A and U (Fig. 5). Reductions in the 
number of setae were arbitrarily omitted from consideration in this analysis, 
though a number of them (e.g., I, P, S, T, V, Y) satisfy the criteria as well as those 
characters that were selected. Knowledge of character HH, the presence or absence 
of endopods on the male’s fifth pair of legs, remains incomplete pending discovery 


VOLUME 96, NUMBER 4 619 


-—Fx<x~xX<0O2=S=CWSCADB 


fm @ 


STS BO Jena as ee 


Fig. 7. Cladogram of hypothesized synapomorphies where criterion in meristic states is relaxed 
to group values with reductions trending similarly. Transformation sequences determined by outgroup 
comparison; further discussion in text. 


of an adult Isaacsicalanus male. The six transformation sequences contain no 
reversals and support the hypothesized phylogenetic relationships shown by this 
cladogram. 

The second cladogram uses transformation sequences in Table 2 that have two 
Or more genera sharing identical derived states; e.g., synapomorphy is proposed 
when two or more genera share the same number of setae on a given structure 
other than the value designated primitive. Nineteen such synapomorphies were 
found, and the resulting hypothesized phylogenetic sequence (Fig. 6) agrees with 
that of Fig. 5. Nine of the characters show one or more reversals, 8 being readily 
accountable by assuming autapomorphic losses of one or more setae by the genus 
showing the reversal. Character D, however, requires autapomorphic losses in 4 
genera. Altering the sequences of Monacilla and Spinocalanus, Spinocalanus and 
Mimocalanus, Mimocalanus and Teneriforma, or Teneriforma and Isaacsicalanus 
in each instance increases the number of reversals measurably. Obviously more 
extensive alterations in the sequence of genera would increase the number of 
reversals still further. 


620 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


The third cladogram (Fig. 7) is based on the synapomorphies proposed for Fig. 
6 and an additional category in which similar trends in loss of setae in a given 
character shown by 2 or more genera were also considered to be synapomorphic. 
This procedure increases the proposed synapomornphies to 24, produces the same 
hypothesized sequence of genera obtained by the two preceding cladograms, and 
decreases the number of reversals to seven. 

The phylogenetic sequence of genera supported by all three analyses appears to 
be a reasonable expression of the data set. Corroboration by additional characters 
to be considered in the future should certainly include a., the sexually modified 
structures of adult males and females, and b., adult pore signatures. 

The implications of these findings include a suggestion of past ages when the 
mixed-layer calanoids may have been taxonomically different from those domi- 
nating at present. Megacalanid-like forms with fully setose oral appendages adapt- 
ed for smaller-particle feeding than extant megacalanids probably occupied at 
least some mixed-layer niches now held by more derived lineages of megacalanoids 
and clausocalanoids. Augaptiloid radiations in the mixed layer may have over- 
lapped with or preceded those of the megacalanids, but the likelihood 1s a changing 
sequence of dominant microplankton feeders in the mixed layer over time. Me- 
gacalanid survivors, spinocalanids, and most of the augaptiloids are now confined 
to meso- and bathypelagic depths and appear to be predaceous or detrital feeders. 

The prospects of developing a relative sequence of taxonomic changes in the 
mixed layer are favorable at this time. The difficulty yet to be faced in the continued 
absence of fossil history of calanoid copepods is the means by which these major 
changes may be related to established geological chronologies. 


Acknowledgments 


It is a pleasure to acknowledge the benefits of David Damkaer’s (1975) excellent 
review of the Spinocalanidae, a work that greatly facilitated phylogenetic analysis 
of spinocalanid genera. For the opportunity to study this unique calanoid I thank 
Ken Smith of Scripps Institution of Oceanography (SIO), the chief Scientist on 
Expedition Oasis and co-worker Roberta Baldwin, SIO, the biological observer 
during ALVIN Dive no. 1217 who detected and sampled the swarm of copepods 
containing J. paucisetus; the paper is contribution no. 4 of Expedition Oasis. The 
final draft was improved by comments from readers ofa previous version, namely, 
Thomas Bowman, Frank Ferrari, Mark Grygier, William Newman, and Richard 
Rosenblatt. This study was funded by the Marine Life Research Program of Scripps 
Institution of Oceanography. 


Literature Cited 


Andronov, V. N. 1973. Taxonomic status of Mecynocera clausi (Copepoda, Calanoida).—Zoolo- 
gicheskii Zhurnal 52(11):1719-1721 [in Russian]. 

Bjérnberg, T. K. S. 1968. Four new species of Megacalanidae (Crustacea: Copepoda).— Antarctic 
Research Series 11:73—90. 

Bowman, T. E. 1976. Miostephos cubrobex, a new genus and species of copepod from an anchialine 
pool in Cuba (Calanoida: Stephidae).— Proceedings of the Biological Society of Washington 
89(1 1):185-190. 

Bowman, T. E., and L. G. Abele. 1982. Classification of the recent Crustacea. Pp. 1-27 in Abele, 


VOLUME 96, NUMBER 4 621 


L. G., ed., The Biology of Crustacea. I. Systematics, the fossil record and biogeography. Aca- 
demic Press, New York and London. , 

Damkaer, D. M. 1975. Calanoid copepods of the genera Spinocalanus and Mimocalanus from the 
central Arctic Ocean, with a review of the Spinocalanidae.— NOAA Technical Report NMFS 
CIRC-391:1-88. 

Eldredge, N., and J. Cracraft. 1980. Phylogenetic patterns and the evolutionary process. Columbia 
University Press, New York, 349 pp. 

Farran, G. P. 1908. Second report on the Copepoda of the Irish Atlantic slope.—Department of 

Agriculture and Technical Instruction for Ireland. Fisheries Branch. Scientific Investigations, 

1906, 2:1-104. 

1936. Copepoda.—British Museum (Natural History), Great Barrier Reef Expedition 1928- 

29. Scientific Reports 5(3):73-142. 

, and W. Vervoort. 195la. Copepoda. Suborder: Calanoida. Family: Spinocalanidae. Genus 

Spinocalanus.—Fiches Identification Zooplancton 39:1—4. 

——., and 1951b. Copepoda. Suborder: Calanoida. Family: Spinocalanidae. Genera Mim- 
ocalanus, Monacilla.—Fiches Identification Zooplancton 40:1-4. 

Frost, B., and A. Fleminger. 1968. A revision of the genus Clausocalanus (Copepoda: Calanoida) 
with remarks on distributional patterns in diagnostic characters.— Bulletin of the Scripps In- 
stitution of Oceanography 12:1—-99. 

Giesbrecht, W. 1888. Elenco dei Copepodi pelagici raccolti dal tenete di vascella Gaetano Chierchia 

durante il viaggio della R. Corvetta “Vettor Pisani” negli anni 1882-1885 e del tenente de 

vascello Francesco Orsini nel Mar Rosso, nel 1884.—R. C. Accademia Nazionale del Lincei, 

Rome (4)4 sem. 2:284-287, 330-338. 

1892. Systematik und Faunistik der pelagischen Copepoden des Golfes von Neapel und der 

angrenzenden Meeresabschnitte.— Fauna und Flora Neapel 19:1-831. 

Grice, G. D. 1971. Deep water calanoid copepods from the Mediterranean Sea. Family Spinocalan- 

idae.— Cahiers de Biologie Marine 12:273-281. 

, and K. Hulsemann. 1965. Abundance, vertical distribution and taxonomy of calanoid co- 

pepods at selected stations in the northeast Atlantic.—Journal of Zoology 146:213-262. 

, and 1967. Bathypelagic calanoid copepods of the western Indian Ocean. — Proceed- 

ings of the United States National Museum 122(3583):1-67. 

Humes, A. G., and M. Dojiri. 1980. A siphonostome copepod associated with a vestimentiferan 
from the Galapagos Rift and the East Pacific Rise.— Proceedings of the Biological Society of 
Washington 93(3):697—707. 

Johnson, R. K. 1982. Fishes of the families Evermannellidae and Scopelarchidae: systematics, 
morphology, interrelationships, and zoogeography.—Fieldiana Zoology, new series, 12:1—252. 

Lonsdale, P. 1977. Clustering of suspension-feeding macrobenthos near abyssal hydrothermal vents 
at oceanic spreading centers.— Deep-Sea Research 24:857-863. 

Park, TaiSoo. 1970. Calanoid copepods from the Caribbean Sea and Gulf of Mexico. 2. New species 
and new records from plankton samples.— Bulletin of Marine Science 20(2):472-546. 

Sanders, R. W. 1981. Cladistic analysis of Agastache (Limiaceae). Pp. 95-114 im Funk, V. A., and 
D. R. Brooks, eds., Advances in Cladistics. 

Sars, G. O. 1903. An account of the Crustacea of Norway. IV. Copepoda, Calanoida.— Bergen, 
171 pp. 

—. 1925. Copépodes particuliérement bathypélagiques provenant des campagnes scientifiques 
du Prince Albert 1°" de Monaco.—Resultats des Campagnes Scientifiques par Albert 1°", Prince 
souverain de Monaco 69:1—408. 

Scott, A. 1909. The Copepoda of the Siboga Expedition. Part 1. Free-swimming, littoral and semi- 
parasitic Copepoda. —Siboga-Expedition, monographie 29a:1-323. 

Sewell, R. B. S. 1947. The free-swimming planktonic Copepoda. Systematic account.—The John 
Murray Expedition 1933-4 8(1):1-303. 

Smith, K. L., Jr., and R. J. Baldwin. Deep-sea respirometry: in situ techniques.—/n Forstner, H., 
and E. Gnaiger, eds., Handbook on Polarographic O, Sensors: Aquatic and Physiological Ap- 
plications. Springer Verlag, Berlin. [in press]. 

Tanaka, O. 1956. The pelagic copepods of the Izu region, middle Japan. Systematic account II. 
Families Paracalanidae and Pseudocalanidae. — Publications of the Seto Marine Biological Lab- 
oratory 5(3):367—406. 


622 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Vervoort, W. 1951. Plankton copepods from the Atlantic sector of the Antarctic. — Verhandelingen 
der Koninklijke Nederlandse Akademie van Wetenschappen, Afd. Natuurkunde. Tweede Sectie, 
Deel 47(4):1-156. 

—. 1957. Copepods from Antarctic and Sub-Antarctic plankton samples.—B.A.N.Z. Antarctic 
Research Expedition, 1929-1931, Reports, Series B (Zoology and Botany) 3:1—160. 

Wheeler, E. H. 1970. Atlantic deep-sea calanoid copeods.—Smithsonian Contributions to Zoology 
(55):1-31. 


Scripps Institution of Oceanography, La Jolla, California 92093. 


PROC. BIOL. SOC. WASH. 
96(4), 1983, pp. 623-631 


EIGHT NEW SPECIES OF INDO-PACIFIC 
CRABS FROM THE COLLECTIONS OF THE 
SMITHSONIAN INSTITUTION 


Tune Sakai 


Abstract.—Four of the species described below were collected during the ex- 
plorations of the U.S. Fish Commission vessel A/batross in 1908 and 1909. The 
remainder are from various sources and were found in the unidentified collections 
of the National Museum of Natural History. The new taxa include representatives 
of eight genera belonging to four families, including the Dorippidae, Leucosiidae, 
Cancridae, and Palicidae. 


Most of the brachyuran crustaceans collected by the Albatross and deposited 
in the Smithsonian Institution were studied by Mary Jane Rathbun, and the 
remaining oxyrhynchous crabs have been studied by Dr. D. J. G. Griffin of the 
Australian Museum. I reexamined the Albatross collections and give herein de- 
scriptions of new species of crabs belonging to the families Dorippidae, Leuco- 
slidae, Cancridae, and Palicidae. In addition, 4 new species of the family Leu- 
cosiidae are described; these were obtained from the Indo-Pacific by other collectors 
on various expeditions. All of these specimens have been deposited in the National 
Museum of Natural History, Smithsonian Institution, Washington, D.C. 


Family Dorippidae MacLeay, 1838 
Ethusa Roux, 1830 


The genus Ethusa comprises about 14 species in the Indo-Pacific, the new 
species is therefore the fifteenth of this genus. In the Philippines, only one species 
of Dorippidae, Ethusina gracilipes Miers (1886), is known. Miers (1886) also 
reported Ethusina sinuatifrons from Japanese seas at a depth of 1875 fathoms, 
but this name is a nomen nudum. 


Ethusa philippinensis, new species 
Fig. la, b 


Material.—1 4, holotype (USNM 195051), 4.5 miles NE of Legaspi Light, 
Lagonoy Gulf, Philippine Islands, 13°12’N, 123°49'18”’E, 72 m, (146 fms), A/- 
batross sta 5453, 7 Jun 1909. 

Description.—The holotype is imperfect and soft-shelled. It is closely related 
to E. hawaiiensis Rathbun (1906; originally E. mascarone hawaiiensis Rathbun), 
the type of which is preserved in the Smithsonian Institution, and which was 
reexamined by the author in 1960. Front of new species produced anteriorly, 
composed of 4 teeth, median incision deepest. Outer orbital tooth triangular, 
situated apparently posterior to front, with its tip directed anteriorly, not laterally. 
Eyes very small. Anterolateral borders moderately swollen posteriorly; in E. ha- 
waiiensis, they are subparallel and only slightly swollen. 


624 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 1. Ethusa philippinensis, new species, male holotype, cl 11.3 mm: a, Dorsal view; b, Anterior 
male pleopod. Ebalia punctulata, new species, female holotype, cl 6.8 mm: c, Carapace; d, Cheliped 
and thoracic leg. 


Dorsal surface of carapace smooth, ““demon-faced” sculpture characteristic of 
Dorippidae faint, but groove around imaginary nose rather distinct. 

Chelipeds slender and symmetrical. Dactyli of ambulatory legs very long, much 
longer than propodi. Anterior pleopod figured in Fig. 1b. 

Measurements.—Length of carapace in the median line 11.3 mm, width 
9.5 mm. 


Leucosiidae Samouelle, 1819 
Ebalia Leach, 1817 
Ebalia punctulata, new species 
ei, le, cl 


Material.—1 2, holotype (USNM 195059), collected at Rongelap Lagoon, west 
of Rongelap Id., 10 m (20 fms), 21 Jun 1946, coll. Tayler. 

Description. —Small species, carapace somewhat rhomboidal. Front moderately 
produced anteriorly, divided into 2 lobes by shallow median incision; anterior 
margins of these lobes straight. Lateral borders markedly convex. Posterolateral 
margins also slightly convex; posterior border transversely straight, lateral angles 
produced into plate-like crest. 

Dorsal surface of carapace punctulate, but anterior third smooth. Surface inside 
hepatic region and on either side of intestinal region moderately depressed. Body 
lacking tubercles and spines. 


VOLUME 96, NUMBER 4 625 


Chelipeds equal, arm tuberculate along anterior and posterior borders, tubercles 
arranged in 2 or 3 longitudinal rows; wrist and palm punctulate. Movable and 
immovable fingers straight, prehensile margins finely denticulate. 

Relationships.— This species is distinctive in having the dorsal surface of the 
carapace minutely punctulate, and in lacking granules and spines on the body. 

Measurements.— Length of carapace 6.8 mm, width 7.1 mm. 


Cryptocnemus Stimpson, 1858 
Cryptocnemus marginatus, new species 
Fig. 2a—c 


Material.—1 6, holotype (USNM 195060), Benkoelen, Sumatra, coll. H. C. 
Kellers, U.S. Navy Eclipse Expedition to Sumatra, Nov 1925. 

Description.—The genus Cryptocnemus is restricted to the Indo-Pacific, con- 
taining about 16 species until now; the new species is therefore the seventeenth. 
The holotype is soft and post-ecdysal. Most thoracic legs are missing and the right 
posterior corner of the carapace is broken and lost. 

Carapace transversely ovoid, dorsal surface flat and smooth, margin elegantly 
bordered with tubercles, regularly and very closely arranged. Front only slightly 
produced beyond outline of carapace, median notch very shallow. Ventral preor- 
bital edge also elegantly bordered with tubercles, its inner anterior notch deep, 
median portion distinctly notched. Outer maxilliped tightly closing the mouth, 
exognath longer and broader than endognath. 

Chelipeds slender, arm, wrist, and palm very swollen, but anterior and posterior 
borders not carinate. Ambulatory legs very slender. 

Male abdomen, and first and second pleopods lost; their characteristics not 
known. 

Measurments.— Length of carapace 6.8 mm, width 7.2 mm. 

Relationships.—This new species is closely related to Cryptocnemus aberrans 
Balss (1938) from Talagi, British Solomon Islands, but in the latter species the 
margin of the carapace is not bordered with tubercles and is strongly upturned. 


Praebebalia Rathbun, 1911 
Praebebalia septemspinosa, new species 
Fig. 2d—h 


Material.—2 6, holotype (USNM 195061) and paratype (USNM 195063), and 
1 2allotype (USNM 195063), NE of island lighthouse, Capitancillo Island, Visayan 
Islands, Philippine Islands, 11°10’N, 124°17'15”E, 90 m, (182 fms), Albatross sta 
5403, 16 Mar 1909. 

Description.—The genus Praebebalia also is restricted to the Indo-Pacific, con- 
taining 4 species until now; the new species is therefore the fifth. Carapace wide 
and subcircular, margin armed with 7 teeth, | pair on either side of lateral margin, 
1 pair at midlength of each posterolateral margin, | pair on either side of posterior 
margin, and finally | on posterior slope of intestinal region. One or 2 small teeth 
on anterior margin of lateral tooth, and | small tooth on hepatic margin. 

Dorsal surface of carapace covered with tubercles of various sizes, those on 
anterior and lateral surfaces large, those on posterolateral and posterior surfaces 


626 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 2. Cryptocnemus marginatus, new species, male holotype, cl 6.8 mm: a, Carapace; b, Cheliped; 
c, First ambulatory leg. Praebebalia septemspinosa, new species, male holotype, cl 13.8 mm: d, Dorsal 
view; e, Abdomen; f, Second male pleopod; g, h, Anterior male pleopod. 


small and indistinct to naked eye. Front produced, composed of 2 teeth, divided 
by median notch, intercalated spine small, external orbital spine larger. 

Chelipeds very long, total length 4.5 times as long as carapace. Arm, wrist, and 
palm finely granulated; fingers inclined about 15 degrees to palm, their prehensile 
edges finely denticulate. Ambulatory legs slender, with no tubercles or granules. 

Male abdomen composed of 5 segments, third to fifth fused, final segment very 
long. Anterior male pleopod slender, distal part furnished with 3 short, 4 median, 
and 4 very long hairs; apical process thin and projected laterally. Second pleopod 
simple (Fig. 2f). 

Measurements.— Length of carapace 13.8 mm, width 14.5 mm, length of che- 
liped 62.0 mm. 

Relationships.— The new species is related to P. extensiva Rathbun (1911), the 
type-species of this genus, but in the latter species, the carapace has only 5 marginal 
spines, and the dorsal surface is smooth, not granulated. 


VOLUME 96, NUMBER 4 627 


Pseudophilyra Miers, 1879 
Pseudophilyra burmensis, new species 
IFS. Sai, [6 


Material.— 1 6, holotype (USNM 195049), Rangoon, Burma, coll. G. E. Gates, 
1928. 

Description.—The genus Pseudophilyra also occurs only in the Indo-Pacific, 
containing 12 species until now; the new species is therefore the thirteenth. 

Carapace almost circular and moderately flattened. Dorsal surface granulate, 
but anterior part of gastric region, postorbital surface, interior of anterolateral 
and central part of intestinal region nongranular and smooth. Front bilobed, each 
lobe rounded and divided by median shallow notch, underlying which, small 
ventral frontal tooth. Intercalated spine small, outer orbital tooth strong and 
obtuse. Eyes very small. Prebuccal plate small and polished. 

Chelipeds strong, arm covered with longitudinally arranged tubercles, distal 
part of arm smooth; wrist and palm smooth except upper and lower margins, 
both slightly granulated. Chelae as long as palm, prehensile edges finely dentic- 
ulate. Ambulatory legs all slender. 

Male abdomen composed of 5 narrow segments, third to fifth coalesced, seventh 
very long and distally narrowed. Anterior male pleopod cylindrical with tip cut 
off obliquely into oval surface, with one margin densely haired, and very long 
hair at base. 

Measurements.— Length of carapace in median line 10 mm, width 10 mm. 

Relationships.—In the arrangement of granules on the upper surface of the 
carapace, the new species is related to Philyra adamsii Bell (1855) but the latter 
species has the front narrower and no median ventral spinule in the median frontal 
sinus, in contrast to the new species. The anterior male pleopod of P. burmensis 
is distinctive (Fig. 3b). 


Leucosia Weber, 1795 
Leucosia bikiniensis, new species 
Fig. 3c-e 


Material.—1 6, holotype (USNM 195050), Sta 3605, 0.5 mi S of west end, 
Bikini Atoll, Marshall Islands, 5 m, (10 fms), 26 Aug 1947, coll. J. P. E. Morrison. 

Description.—Carapace very broad, breadth exceeding length. Front produced 
slightly anteriorly, divided into 2 lobes by median shallow sinus. Hepatic border 
produced into low triangular lobe. Lateral border densely furnished with hairs; 
narrow crest present along posterolateral and posterior borders. Dorsal surface 
smooth, coloration faded; ocelli or stripes usually found in Leucosia not observed. 

Chelipeds rather short and robust. Left cheliped, arm bearing 2 very large 
tubercles followed by | basal and | smaller distal tubercle on anterodorsal border; 
cluster of about 4 small tubercles at upper basal surface; about 6 medium-sized 
tubercles on posterior border; and finally, 5 smaller tubercles in cluster at posterior 
basal surface, covered with velvety tomentum. 

Right cheliped, arm bearing 3 large tubercles on dorsoanterior border; otherwise 
arrangement of smaller tubercles about same as those of left cheliped. 

Wrist, palm, and fingers carinate along inner and outer borders. Outer distal 


628 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


part of wrist bearing row of several small tubercles traversed by row of few 
tubercles; basal end of immovable finger armed with 2 oblique rows of small 
tubercles. 

Male abdomen like that of related species, composed of 4 pieces, second, and 
third broad, third segment armed with spinule in middle. Anterior male pleopod 
thin, distal end thickened and divided into 2 hemispheres, covered with hairs. 
Inner hemispheres armed with low spine, outer hemisphere slightly higher, with- 
out spine. 

Measurements.— Length of carapace 7.0 mm, width 7.3 mm. 

Relationships.— This species is closely related to Leucosia insularis Takeda and 
Kurata (1976), from the Bonin Islands (=Ogasawara Islands), but in the latter 
species the thoracic sinus is marked with one large and several smaller tubercles 
instead of 2 ridges of small tubercles and a few larger tubercles between the 2 
ridges in L. bikiniensis. In the Bonin species, the apex of the male pleopod is 
globular, while in the new species it is divided into two hemispheres. 


Cancridae Latreille, 1803 
Cancer Linnaeus, 1758 


Cancer is the oldest and fundamental genus of the decapod crustaceans, estab- 
lished by Linnaeus in 1758. In the Indo-Pacific, this genus comprises 8 species 
until now: 


Cancer gibbosulus (De Haan, 1835) 

= Trichocarcinus affinis Miers, 1879 

Cancer novaezelandae (Jacquinot, 1853) 

Cancer japonicus Ortmann, 1893 

Cancer amphioetus Rathbun, 1898 

= Trichocarcinus dentatus Miers, 1879 (preoccupied) 
Not Cancer dentatus Herbst, 1785 
Not Cancer dentatus Bell, 1835 

= Cancer bullatus Balss, 1922 

Cancer tumifrons Yokoya, 1933 

Cancer nadaensis Sakai, 1969 

Cancer sakaii Takeda & Miyaki, 1972 

= Platepistoma anaglyptum Balss, 1922 (preoccupied) 

Cancer guezei Crosnier, 1976 


Cancer luzonensis, new species 
Fig. 3f 


Material.—1 2, holotype (USNM 195064), off Matocot Point, Luzon Island, 
Philippine Islands, 13°41'50”N, 120°58’30’E, 86 m, (172 fms), Albatross sta 5289, 
22 Jul 1907. 

Description. — This species has 3 frontal teeth, median very small and in a lower 
ventral position. Orbital teeth strong. Anterolateral margin divided into 9 teeth, 
subequal in size and shape and arranged in arch, last tooth situated at outermost 
corner. Posterolateral border slightly concave, armed with about 8 teeth behind 
last anterolateral tooth. Posterior margin also slightly concave, smooth, without 
teeth: 


VOLUME 96, NUMBER 4 629 


Fig. 3. Pseudophilyra burmaensis, new species, male holotype, cl 10 mm: a, Dorsal view; b, Anterior 
male pleopod. Leucosia bikiniensis, new species, male holotype, cl 7.0 mm: c, Carapace and chelipeds; 
d, Thoracic sinus; e, Anterior male pleopod. Cancer luzonensis, new species, female holotype, cl 34.0 
mm: f, Dorsal view. Palicus bidentatus, new species, female holotype, cl 7.1 mm: g, Carapace; h, 
Cheliped; i, Second ambulatory leg; j, Third ambulatory leg. 


Dorsal surface of carapace deeply areolate, grooves separating gastric, cardiac, 
and branchial regions deep and distinct. These regions divided into subregions; 
anterior surface of gastric region and lateral surfaces of hepatic and branchial 
regions sparingly tuberculate. Cardiac and intestinal regions also indistinctly tu- 
berculate. Chelipeds subequal in size and shape; arm short and almost hidden 


630 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


beneath anterolateral margin, upper surface smooth. Upper surface of wrist smooth, 
anterior margin granulate and haired. Outer surface of palm marked with 5 lon- 
gitudinal rows of granules, distal portions of these rows marked with large tu- 
bercles; outer surface marked with 4 longitudinal rows of granules. Fingers lightly 
pigmented with brown, prehensile margins armed with 4 or 5 teeth, proximal 1 
-or 2 large. Ambulatory legs rather slender, anterior edges of carpus and propodus 
spinulate and haired. 

Measurements.— Length of carapace measured between the tips of the frontal 
teeth and the posterior margin 34 mm, width of same measured between the tips 
of the anterolateral teeth 47 mm. 

Relationships.—This species is related to Cancer nadaensis Sakai (1969) in its 
general aspect, but the dorsal surface of the carapace is deeply areolated, differing 
from the smooth surface of C. nadaensis, and the last anterolateral tooth is less 
produced laterally. 


Palicidae Rathbun, 1898 
Palicus Philippi, 1838 
Palicus bidentatus, new species 
Fig. 3g-] 


Material.—1 2, holotype (USNM 195065), SE of Hong Kong Island, 21°42'N, 
114°50’E, 19 m, (38 fms), Albatross sta 5302, 9 Aug 1908. 

Description.—This specimen is very old and fragmentary, the chelipeds and 
ambulatory legs have fallen off. Carapace broadly rectangular. Front not much 
produced anteriorly, composed of 4 teeth; upper orbital teeth also 4 in number, 
each separated by fissure. Upper surfaces of these teeth sparsely granulate, pos- 
terior to these teeth surface of anterior carapace transversely depressed. Around 
eyestalk, ventral ridge of orbit armed with 1 outer and 2 inner teeth; these teeth 
all strong and high compared with those of other species. 

Hepatic border very short; branchial border armed with 2 obtuse densely gran- 
ulate processes at lateral border. Posterolateral and posterior borders united, trans- 
verse, without teeth or spines. 

Dorsal surface of carapace strongly uneven, sparingly covered with granules. 
Regions of carapace not clearly separated, but each with patches of thick granules. 
Gastric region with 3 rhomboidal patches of granules, cardiac region divided into 
2 by median groove, intestinal region vaguely separated into 4. Between cardiac 
and intestinal regions, surface deeply and transversely depressed. Inner surfaces 
of the anterolateral and posterolateral borders each with patch of thick granules. 

Chelipeds slender; arm, wrist, and palm not markedly granulate; prehensile 
edges entire, blade-like. Third ambulatory leg largest, second and first pairs little 
smaller, fourth pair very small and filiform. In first 3 ambulatory legs, merus 
thickened proximally, subdistal portion thin, distal outer part armed with strong 
tooth; carpus, propodus and dactylus with long hairs along anterior border. 

Measurements.— Length of carapace 7.1 mm, width 10.5 mm. 

Relationships.— Among the numerous species of Palicus, this species is peculiar 
in having the carapace transversely rectangular and its anterolateral border is 
armed with 2 obtuse processes, covered with granules. Around the orbit, this 
species has 3 dorsal, 1 lateral, 1 outer ventral, and 2 inner ventral teeth. 


VOLUME 96, NUMBER 4 631 


Acknowledgments 


I wish to express my deep gratitude to Dr. Fenner A. Chace, Jr., who originally 
made the specimens available for study. Thomas E. Bowman and Raymond B. 
Manning edited the final drafts of the manuscript. 


Literature Cited 


Balss, H. 1938. Die Dekapoda Brachyura von Dr. Sixten Bocks Pazifik-Expedition 1917-1918.— 
GGteborgs Kungliga Vetenskaps-och Vitterhets-Samhilles Handlingar, Femte Féljenden, series 
b 5(7):1-85. 

Bell, T. 1855. A monograph of the Leucosiadae, with observations on the relations, structure, habits 
and distribution of the family; a revision of the generic characters; and descriptions of new 
genera and species. Hora Carcinologicae, or Notices of Crustacea, I.—Transactions of the 
Linnean Society 21:277-314. 

Miers, E. J. 1886. Report on the Brachyura collected by H.M.S. Challenger during the years 1873- 
1876.—Report on the Scientific Results of the Voyage of H.M.S. Challenger during the years 
1873-1876, Zoology, 17:xli + 362. 

Rathbun, M. J. 1906. The Brachyura and Macrura of the Hawaiian Islands.— Bulletin of the U.S. 

Fish Commission for 1903, part 3:827—930. 

1911. Marine Brachyura. The Percy Sladen Trust Expedition to the Indian Ocean in 1905, 
3(9).— Transaction of the Linnean Society, London, Zoology, (2)14(2):191—261, pls. 15-20. 
Sakai, T. 1969. Two new genera and twenty-two new species of crabs from Japan.— Proceedings of 

the Biological Society of Washington 82:243-280. 

Takeda, M., and Y. Kurata. 1976. Some species collected by coral fishing boats. Crabs of the 
Ogasawara Islands, IIIJ.— Bulletin of the National Science Museum, Tokyo, series A (Zoology) 
2(1):19-32. 


562 Jyomyoji-machi, Kamakura, Kanagawa-ken, Japan. 


PROC. BIOL. SOC. WASH. 
96(4), 1983, pp. 632-637 


A NEW SPECIES OF THE SERRANID FISH GENUS 
PLECTRANTHIAS (PISCES: PERCIFORMES) FROM 
THE SOUTHEASTERN PACIFIC OCEAN, WITH 
COMMENTS ON THE GENUS ELLERKELDIA 


Phillip C. Heemstra and William D. Anderson, Jr. 


Abstract.—Plectranthias exsul, new species, is described from six specimens 
collected off the Juan Fernandez Islands and from the Nazca Ridge off the coast 
of Chile. This new species is the first Plectranthias to be reported from the eastern 
Pacific Ocean. The distinction between Plectranthias and Ellerkeldia is discussed. 


For several years we have been working on a revision of the American Anthiinae. 
Among the considerable material examined are six specimens of a new species 
of Plectranthias Bleeker, 1873, from the southeastern Pacific. Rather than delay 
description of this new species until the completion of our revision, we decided 
to publish the description separately. 

Type-specimens are deposited in the Academy of Natural Sciences of Phila- 
delphia (ANSP); Bernice P. Bishop Museum, Honolulu (BPBM); Museum of 
Comparative Zoology, Harvard University (MCZ); National Museum of Natural 
History, Smithsonian Institution (USNM); and Zoological Museum, Moscow State 
University (ZMMU). Measurements and counts were made following Anderson 
and Heemstra (1980). 


Plectranthias exsul, new species 
Figs. 1-2 


Plectranthias sp., Parin et al., 1981:14 (brief description of Nazca Ridge speci- 
mens). 


Holotype.— ANSP 127843, 158 mm SL (standard length); Juan Fernandez is- 
lands (33°37'S, 78°49'W); Feb 1957; R. Manning, collector. 

Paratypes.—MCZ 52520, 2 specimens 134 & 140 mm SL; Juan Fernandez 
islands; 140 to 165 m; 27 Jan 1966; R/V Anton Brunn Cruise 13, Stn. 35; 40’ 
otter trawl. USNM 176577, 158 mm SL; Juan Fernandez islands (33°38’S, 
78°50’W); 31 Mar 1945. BPBM 27978, 133 mm SL; Nazca Ridge (25°45’S, 
85°29'W); 200 to 225 m; 1 Nov 1979; R/V Ikhtiandr Trawl 59. ZMMU P-16022, 
150 mm SL; data as for BPBM 27978. 

Diagnosis.—Dorsal-fin rays X, 15 or X, 16. Anal-fin rays III, 7. Pectoral-fin 
rays 16 or 17 (usually 16); dorsalmost (and occasionally ventralmost) ray un- 
branched, the rest branched. Branched caudal-fin rays 8 + 7. Lateral-line scales 
40 to 46; scales between middle of spinous dorsal fin and lateral line 1'2 to 2%. 
Lateral part of snout and anterior part of lower jaw naked. Maxilla naked or with 
very few scales. Small splint-like supramaxilla usually present. Preopercle serrate, 
with or without antrorse spine on lower limb. Gillrakers 8 to 10 + 18 to 21, total 
26 to 31. Second soft ray of dorsal fin and one of dorsalmost branched rays of 


VOLUME 96, NUMBER 4 633 


Fig. 1. Plectranthias exsul, holotype, ANSP 127843, 158 mm SL. 


caudal fin elongate. Posterior margin of caudal fin truncate to slightly concave. 
Vertebrae 10 + 16. Arrangement of predorsal bones, anterior neural spines, and 
anterior pterygiophores of dorsal fin 0/0 + 0/2/1 + 1/1/ (using notation of Ahlstrom 
et al. 1976). 

Description. — Morphometric data are given in Table |. Dorsal fin continuous, 
not divided to base between spinous-and soft-rayed parts; fourth or fifth spine of 
dorsal fin longest, 1.6 to 2.0 times length of last spine of dorsal fin; second soft 
ray of dorsal fin elongate, longer than longest spine of dorsal fin. Anal fin margin 
subangular to slightly pointed; second spine of anal fin about twice length of first, 
much stouter than third; third shorter than second. Pectoral fin pointed; middle 
rays longest, usually reaching to vertical through base of third spine of anal fin. 
Pelvic fin reaching or falling well short of anus. Principal caudal-fin rays 9 + 8; 
procurrent caudal-fin rays 9 or 10 dorsally, 7 to 9 ventrally. 


Body depth 2.5 to 3.0, head length 2.5 to 2.7 in SL. Orbit 3.8 to 4.5, interorbital 
width 5.8 to 6.7 in head length. Maxilla reaching or almost reaching vertical 
through posterior border of orbit; posteroventral corner of maxilla with rounded 
projection. Anterior naris oblique, at distal end of short tube (posterior border of 
tube highest); posterior naris elliptical to subcircular. Scales ctenoid. Scales on 
dorsum of snout extending anteriorly on each side of naked mid-dorsal area nearly 
to upper lip (in one specimen anterior part of mid-dorsal area covered with scales). 
Interorbital region, cheek, preopercle, opercle, subopercle, interopercle, and pos- 
terior 13 to % of ventral surface of lower jaw with scales. Scales on cheek extending 


634 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Table 1.—Morphometric data for Plectranthias exsul. The holotype is denoted by an asterisk. 
Standard length is in mm; other measurements, in percentage of standard length. 


BPBM ZMMU ANSP USNM 
Measurement 27978 MCZ 52520 P-16022 127843* 176577 

Standard length 133 134 140 150 158 158 
Head, length 37.7 38.4 39.1 38.2 37.6 37.8 
Snout, length 10.6 9.3 10.0 10.3 9.4 9.8 
Orbit, diameter 9.5 9.9 10.2 8.5 9.5 8.4 
Postorbital length of head 72 20.0 18.8 18.5 18.6 19.4 
Upper jaw, length 18.3 19.6 19.6 75) 19.0 18.7 
Maxilla, width 52) 6.3 6.2 5.0 Sa 5.5 
Interorbital width 6.0 6.3 5.9 6.1 6.5 6.3 
Body, depth 33.7 37.3 39.6 36.1 35.2 36.5 
Predorsal length 35.8 39.9 39.3 37.0 37.9 39.5 
Preanal length 70.6 64.9 66.4 70.1 64.7 66.2 
Caudal peduncle, length 21.2 20.9 20.4 22.1 DDT 22.3 
Caudal peduncle, depth 10.3 12.4 13.1 10.7 11.2 10.7 
Pectoral fin, length ca. 31.7 33.3 30.7 33.3 33.2 32.5 
Pelvic fin, length 229 26.1 Ds ca. 22.3 23.4 24.1 
Anal fin, length 28.0 29.9 28.9 26.5 ca. 30.8 26.8 
Upper caudal-fin lobe, length > 30.4 28.2 28.6 ca.24.5 ca. 29.9 ~ 
Lower caudal-fin lobe, length DAD 23.9 26.1 ca. 23.1 ca. 24.0 — 
Third dorsal spine, length 57 15.9 16.3 >14.8 >16.8 >15.4 
Fourth dorsal spine, length 18.2 18.7 18.4 >16.7 >18.8 >16.1 
Longest dorsal spine, length 18.2 18.7 18.4 >16.7 19.9 18.1 
(4th) (4th) (4th, Sth) (4th) (Sth) (Sth) 
First anal spine, length 5 9.3 8.4 8.7 9.8 8.9 
Second anal spine, length 19.0 17.3 — 16.1 18.7 16.6 
Third anal spine, length >15.3 14.9 15.9 14.7 3) 7/ 14.7 


anteriorly to vertical through middle of orbit. Branchiostegal rays and membranes, 
and gular area naked (few scales anteriorly along midline of gular area in some 
specimens). Opercle with 3 spines; dorsalmost blunt and inconspicuous, middle 
one largest. Few to numerous small to well-developed serrae and/or irregularities 
on distal margins of subopercle and interopercle at or near their junction. Bran- 
chiostegal rays 7; anterior 3 inserting along ventral edge of hyoid arch, posterior 
4 inserting laterally on arch. Longest gillrakers slightly longer than longest gill 
filaments. Pseudobranch well developed, with 23 to 31 filaments. 

Proximal half of soft dorsal and anal fins scaly; pectoral, pelvic and caudal fins 
scaly basally; no enlarged axillary scales at base of pelvic fin. Rows of cheek scales 
8 to 12. Series of circum-peduncle scales 18 to 22. No smaller auxiliary scales at 
bases of body scales. Lateral line complete, extending to base of caudal fin; running 
a few scale rows below dorsal fin parallel to dorsal body contour, curving to mid- 
lateral axis of body somewhat posterior to vertical through posterior end of dorsal- 
fin base. Lateral-line tubes simple. Scales from dorsal-fin origin to lateral line 4 
or 5; scales from lateral line to anal-fin origin 13 to 17. 

Upper jaw with band of very small depressible teeth; band broadest in anterior 
portion of jaw where inner teeth are variously enlarged into posteriorly directed 
conical and caniniform teeth; 1 to 3 stout exserted canines on each side near 
symphysial diastema. Lower jaw with narrow band of small depressible conical 


VOLUME 96, NUMBER 4 635 


Fig. 2. Plectranthias exsul, paratype, MCZ 52520, 140 mm SL. 


teeth; band broadened at anterior end of jaw; | to 3 stout exserted canines on 
each side of jaw near symphysial diastema; | to 3 stout recurved canines on each 
side of jaw about one-third distance from symphysis to posterior termination of 
dentition; inner teeth of band somewhat enlarged; inner teeth near symphysis 
recurved and distinctly enlarged. Vomer and palatines with cardiform teeth; vo- 
merine teeth in chevron-shaped patch; palatine teeth in band, anterior portion of 
band somewhat broadened and usually slightly curved toward vomer. No teeth 
on tongue or pterygoids. 

Eleventh and twelfth vertebrae with ventrolateral foramina. Epipleural ribs on 
anterior 12 or 13 vertebrae; pleural ribs on vertebrae 3 through 10. No hypural 
fusions. Trisegmental pterygiophores associated with anal fin 1 or 2, with dorsal 
fin none. 

Color.—In alcohol, body and head uniformly straw colored; color in life un- 
known. 

Comparisons. —Plectranthias exsul has more lateral-line scales than any other 
species in the genus except P. tay/ori Randall, 1980; however, it is separated easily 
from P. taylori in having more gillrakers (total on first gill arch 26 to 31 vs. 17 
or 18), fewer dorsal soft rays (15 or 16 vs. 18), and more pectoral-fin rays (16 or 
17 vs. 14). 

Plectranthias exsul is also similar to P. kelloggi (Jordan & Evermann, 1903), 
with which it shares the following characters: second soft ray of dorsal fin and 
one of dorsalmost branched rays of caudal fin elongated, similar fin-ray counts, 
and similar dentition. In addition to having more lateral-line scales (40 to 46 vs. 
32 to 38), P. exsul differs from P. kelloggi in having more gillrakers (total on first 


636 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


gill arch 26 to 31 vs. 20 to 24) and in possessing a naked or almost naked maxilla 
and a partially naked mandible (anterior one-third to two-thirds without scales) 
vs. a partially scaly maxilla and a scaly mandible. 

Distribution. — Plectranthias exsul is known only from off the Juan Fernandez 
islands and from the Nazca Ridge off the coast of Chile. Depths of capture range 
from 140 to 225 meters. 

Etymology.—The Latin noun exsu/ (meaning “‘exile’’) is used in allusion to the 
eastern Pacific distribution of this species; P. exsul is the first species of Plec- 
tranthias known from the eastern Pacific region. 

Remarks.—In his revision of the genus Plectranthias, Randall (1980) recognized 
30 species, 13 of which he described as new. Five other new species have recently 
been placed in Plectranthias by Fourmanoir and Rivaton (1980), Katayama and 
Masuda (1980), Fourmanoir (1982), and Raj and Seeto (1983). It should be 
pointed out that our assignment of P. exsul to Plectranthias is based on Randall’s 
definition of the genus. In order to precisely place P. exsu/ or any other species 
currently assigned to the genus, more study, particularly of internal morphology, 
is needed. It seems likely that a reassessment of currently available data coupled 
with an analysis of new information will lead to a modification of the description 
of Plectranthias provided by Randall (1980) and to the recognition at the generic 
level of one or more of the eight genera subsumed by him into Plectranthias. 

Although considerations of the limits and the diagnostic characteristics of the 
genus Plectranthias are beyond the scope of this study, we feel that a comment 
on the generic-level classification proposed by Randall is in order. Randall (1980: 
102) considered the genus Ellerkeldia Whitley, 1927, to be “closely related to 
Plectranthias.”’ He distinguished the two genera by the “smaller scales (40 to 50 
in the lateral-line series .. .)”’ and “‘the configuration of the head” being ““more 
sharply linear” in Ellerkeldia. Nevertheless Plectranthias taylori Randall, 1980, 
has 40 to 41 lateral-line scales, our new species, P. exsul, has 40 to 46, and E. 
Jamesoni (Ogilby, 1908) has 38 to 42. Judging by the available illustrations, there 
are several species of Plectranthias with head profiles as “sharply linear” as those 
of species of Ellerkeldia. Although neither of Randall’s criteria will serve to 
distinguish these two genera, it appears that number of vertebrae is useful. Each 
of the six species of E//erkeldia we examined has 27 vertebrae (10 + 17), whereas 
all species of Plectranthias, for which counts are available (Katayama and Masuda 
1980; Randall 1980; and the present work), have 26. Provisionally then, we accept 
Randall’s definition of the genus Plectranthias and consider it distinct from the 
genus Ellerkeldia, but are well aware that neither genus is satisfactorily defined 
or differentiated from related genera. 


Acknowledgments 


The curators of fishes at the Academy of Natural Sciences of Philadelphia, 
Museum of Comparative Zoology, and the National Museum of Natural History 
allowed us to examine specimens in their care. J. E. Randall forwarded material 
he received from the P. P. Shirshov Institute of Oceanology in Moscow; we are 
grateful to N. V. Parin of that Institute for permission to deposit a paratype in 
the Bernice P. Bishop Museum. J. L. Russo made the photograph for Fig. 1. Susan 
Jewett, J. F. McKinney, and G. C. Van Dyke provided radiographs. G. D. Johnson 


VOLUME 96, NUMBER 4 637 


reviewed the manuscript. This paper is contribution number 58 of the Grice 
Marine Biological Laboratory. This research was supported in part by National 
Science Foundation Grant GB-40063 to P. C. Heemstra. 


Literature Cited 


Ahlstrom, E. H., J. L. Butler, and B. Y. Sumida. 1976. Pelagic stromateoid fishes (Pisces, Perciformes) 
of the eastern Pacific: kinds, distributions, and early life histories and observations on five of 
these from the northwest Atlantic.— Bulletin of Marine Science 26:285-402. 

Anderson, W. D., Jr., and P. C. Heemstra. 1980. Two new species of western Atlantic Anthias (Pisces: 
Serranidae), redescription of A. asperilinguis and review of Holanthias martinicensis. —Copeia 
1980:72-87. 

Bleeker, P. 1873. Sur les espéces indo-archipelagiques d’Odontanthias et de Pseudopriacanthus. — 
Nederlandsch Tijdschrift voor de Dierkinde 4:235-240. 

Fourmanoir, P. 1982. Trois nouvelles espéces de Serranidae des Philippines et de la Mér du Corail 
Plectranthias maculatus, Plectranthias barroi, Chelidoperca lecromi.—Cybium 6:57-64. 
Fourmanoir, P., and J. Rivaton. 1980. Plectranthias randalli n. sp., un nouveau serranidé (anthiiné) 

du sud-ouest Pacifique.— Revue francaise d’Aquariologie 7:27-28. 

Jordan, D. S., and B. W. Evermann. 1903. Descriptions of new genera and species of fishes from 
the Hawaiian Islands.— Bulletin of the United States Fish Commission 22:161—208. 

Katayama, M., and H. Masuda. 1980. Two new anthiine fishes from Sagami Bay, Japan.—Japanese 
Journal of Ichthyology 27:185-190. 

Ogilby, J. D. 1908. On new genera and species of fishes.—Proceedings of the Royal Society of 
Queensland 21:1—26. 

Parin, N. V., G. A. Golovan, N. P. Pakhorukov, Yu. I. Sazonov, and Yu. N. Shcherbachev. 1981. 
Fishes from the Nazca and Sala-y-Gomez underwater ridges collected in cruise of R/V “Ikh- 
tiandr.” In: Fishes of the Open Ocean.—Institute of Oceanology, Academy of Sciences of the 
USSR, Moscow (“1980”). pp. 5-18. 

Raj, U., and J. Seeto. 1983. A new species of the Anthiine fish genus Plectranthias (Serranidae) from 
the Fiji Islands.— Japanese Journal of Ichthyology 30:15-17. 

Randall, J.E. 1980. Revision of the fish genus Plectranthias (Serranidae: Anthiinae) with descriptions 
of 13 new species.— Micronesica 16:101-187. 

Whitley, G. P. 1927. Studies in ichthyology. No. 1.—Records of the Australian Museum 15:289- 
304. 


(PCH) J. L. B. Smith Institute of Ichthyology, Private Bag 1015, Grahamstown 
6140, Republic of South Africa; (WDA) Grice Marine Biological Laboratory, 
College of Charleston, Charleston, South Carolina 29412. 


PROC. BIOL. SOC. WASH. 
96(4), 1983, pp. 638-644 


REDESCRIPTION OF THE BRAZILIAN LABRISOMID 
FISH STARKSIA BRASILIENSIS 


J. T. Williams and A. M. Smart 


Abstract.— Starksia brasiliensis (Gilbert), known only from southern Brazil, is 
redescribed based on 16 specimens collected from near Sao Paulo, Brazil, and all 
S. brasiliensis specimens mentioned in earlier literature. Starksia brasiliensis is 
distinguished from other species in the S. oce/lata complex by a combination of 
the following characters: narrow, pale Y-shaped mark on cheek; ring-shaped marks 
on cheek bright red in life; no bars on lips; typically 16 (range 15-17) scales in 
arched part of lateral line; and typically 7 (range 7—9) total gill rakers on the first 
arch. 

The known geographic range of Starksia brasiliensis is extended from Maceio 
and Salvador, Brazil, some 1600 km south to Sao Paulo, Brazil. 


Starksia brasiliensis, a poorly known labrisomid (sensu George and Springer 
1980) previously represented by only five specimens, inhabits rocky, coralline 
areas in the shallow coastal waters of southern Brazil. Sixteen specimens of S. 
brasiliensis, collected by one of us (AMS) near Sao Paulo, allow an expanded 
analysis of character variation. The variation reported herein significantly in- 
creases the ranges for meristic characters given by Greenfield (1979), and has 
made it necessary to redescribe the species. These specimens also extend the 
geographic range of the species southward to Sao Paulo, Brazil. 

The taxonomic validity of Starksia brasiliensis (Gilbert, 1900) has been in doubt 
since it was described. Longley and Hildebrand (1941) placed it in the synonymy 
of S. ocellata (Steindachner, 1876). They did not give reasons for taking this 
action, although they did examine one specimen of S. brasiliensis collected off 
the coast of Salvador, Brazil. BOhlke and Springer (1961) reviewed Starksia and 
commented on the low anal-fin ray and scale counts of a paratype of S. brasiliensis. 
Because there was so little material available, they chose to retain S. brasiliensis 
in the synonymy of S. ocellata. In his review of the S. ocellata complex, Greenfield 
(1979) examined four specimens (including both types) from southern Brazil and 
gave this population species status based on three major characters: a narrow, 
pale Y-shaped mark on the cheek, no bars on the lips, and a single row of in- 
fraorbital pores. We find the first two characters useful for species recognition but 
the infraorbital pores are frequently in a double row, thus limiting the usefulness 
of these pores as a distinguishing character. 

Counts and measurements follow Greenfield (1979). In addition, tooth counts 
refer only to the teeth in the outermost row. The cephalic sensory pore series are 
delimited as shown in Fig. 1 (counts include all pores in each series). Predorsal, 
preanal, and prepelvic lengths refer to the distance from the snout tip to the 
anterior of the base of the respective fin. Procurrent caudal-fin rays are those 
caudal elements with less than two segments. 

The following institutional abbreviations are used: BMNH—British Museum 
(Natural History); CAS-SU—Stanford University collection at the California 
Academy of Sciences; GCRL— Gulf Coast Research Laboratory in Ocean Springs, 


VOLUME 96, NUMBER 4 639 


AN 


ec 
no) S==as8as==9 
— (ie Viera Se 
. A 


POP 


Fig. 1. Semidiagrammatic illustration of cephalic sensory pore series. Lines indicate first and last 
pores in each series. Abbreviations: AN, anterior nostril; IO, infraorbital series; LT, lateral temporal 
series; MD, mandibular series; PN, posterior nostril; POP, preopercular series; SO, supraorbital series; 
ST, supratemporal series. 


Mississippi; MZUSP— Museu de Zoologia da Universidade de Sao Paulo; UF— 
Florida State Museum. Other abbreviations: IO, infraorbital pore series; LT, 
lateral temporal pore series; MD, mandibular pore series; POP, preopercular pore 
series; SO, supraorbital pore series; ST, supratemporal pore series. 


Starksia brasiliensis (Gilbert, 1900) 
Fig. 2 


Brannerella brasiliensis Gilbert, 1900:180, pl. 9, fig. 1 (holotype CAS-SU 7750; 
type locality: coral reef near Macei0, Brazil). 


640 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Diagnosis.— Distinguished from other species in the Starksia ocellata complex 
by the following combination of characters: cheek with very narrow, pale Y-shaped 
mark; ring-shaped marks on cheek bright red in life; pigment on lips restricted 
to tips of jaws; typically 16 scales in arched part of lateral line (range 15-17); 
typically 7 total gill rakers on first arch (range 7-9). 

Description.— Ranges for selected meristic characters are as follows: dorsal fin 
XX-XXI, 7-9 (mode XXI, 8); total dorsal elements 28-30 (mode 29); anal fin 
II, 16-18 (mode 17, spines not connected in males); pectoral fin 13-14 (mode 
14); pelvic fin I, 2 apparent (a third rudimentary ray observable in some radio- 
graphs); caudal-fin rays 7 on dorsal hypurals, 6 on ventral hypural plate, none 
branched; procurrent dorsal caudal-fin rays 5—6 (mode 6); procurrent ventral 
caudal rays 4—6 (mode 5); precaudal vertebrae (those without distinct hemal spine) 
10; caudal vertebrae (those with distinct hemal spine, plus the complex hypural 
bearing centrum) 23-25 (mode 24, CAS-SU 53510 paratype with 20 is anoma- 
lous); scales in arched part of lateral line 15—17 (mode 16), scales in straight part 
19-22 (mode 20); gill rakers on first arch 2—3+1+4—6 (mode 2+1+4); last 
pleural rib on vertebra 10; last epipleural rib difficult to discern, but approximately 
on vertebra 1 7—19 (discernable in radiographs of only 3 specimens); pseudobran- 
chial filaments 3—5 (mode 4); premaxillary teeth 46-57 (13 specimens); dentary 
teeth 51-73 (8 specimens); vomerine teeth 8-14 (mode 12); palatine teeth (left— 
right) 3-2 to 5-5 (mode 4-4). Cephalic sensory pores (Fig. 1, range followed by 
mode in parentheses): ST 12-13 (13), LT 8-11 (9), POP 7-11 (9), SO 6-10 (7), 
IO 8-13 (11-12), MD S. 

Upper and lower jaws each with outer rows of stout conical teeth enclosing 
band of small conical teeth near symphyses. Vomer and palatines with row of 
stout conical teeth, sometimes flanked posteriorly, or mesially, respectively, by 
small conical teeth. Small, conical upper pharyngeal teeth in rounded patch of 
about 3 irregular rows. 

Pigmentation of preserved specimens as described by Greenfield (1979). Life 
colors taken from kodachrome slides of freshly collected specimens as follows: 
iris of eye yellow; bright red ring-like marks on cheeks, two on base of each 
pectoral fin, and one at upper end of gill opening. Red spots scattered over dorsal, 
anal, and caudal fins. Other markings the same as in preserved specimens. 

Of the 21 known specimens of Starksia brasiliensis, there are 12 females and 
nine males. The largest female is 34.1 mm SL and the largest male is 30.9 
foovoal SL, 

The data for the Sao Paulo population (available on request) were analyzed for 
sexual differences using a t-test, but no statistically significant sexual differences 
were found for any of the characters examined. The northern and southern pop- 
ulations have mean numbers of segmented dorsal-fin rays (northern 7.4 vs. south- 
ern 8.1), segmented anal-fin rays (16.8 vs. 17.7), and total vertebrae (33.0 vs. 
33.9) that are significantly different as indicated by t-tests (dorsal rays P < 0.001, 
t = 6.182; anal rays P < 0.001, t = 7.877; vertebrae P < 0.001, t = 7.728). These 
differences may represent clinal variation, but this cannot be confirmed until 
specimens from geographically intermediate areas become available. 

Distribution. — Starksia brasiliensis is known from Macei0, south to Sao Paulo, 
Brazil, in depths to 13 m. One specimen collected from a cable was reported from 
61 m (Longley and Hildebrand 1941; label with specimen gives 55-62 m), but 


-VOLUME 96, NUMBER 4 641 


Fig. 2. Starksia brasiliensis. A. Male, 27.2 mm SL (MZUSP uncat.). B. Female, 31.8 mm SL 
(MZUSP uncat.). 


we are hesitant to consider this depth within the normal range for the species 
(reasons discussed below). 

The Sao Paulo specimens represent a southward range extension of approxi- 
mately 1600 km to 23°44'S. We believe that the apparent disjunction in the species’ 
distributional range reflects the limited collecting efforts in the intervening area 
and that the species will be commonly found in rocky and coral bottom habitats 
in that area. 

The southern Brazil collecting sites were along the leeward shores of two steeply 
sloping, heavily forested islands. Large boulders (up to 5 x 5 x 2 m) covered the 
slopes above and below water, with those underwater covered with algae, bryo- 
zoans, sponges, small corals and other encrusting organisms. Small shrimp, lob- 
sters, crabs, snails, and nudibranchs were observed on the encrusting growth. 

Remarks.—The paratype (CAS-SU 53510) of Starksia brasiliensis has caused 
confusion regarding the range of certain counts. BOhlke and Springer (1961) com- 
mented on the low anal-fin ray and scale counts, and Greenfield (1979) gave an 
unusually low vertebral count for this specimen. Close examination of radiographs 


642 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


of this specimen reveals that two pairs of centra, 6-7 and 14-15, are almost 
completely fused, each pair appearing as only one centrum. Nonetheless, the 
complex nature of the centra is evident since each bears two neural spines and 
two hemal arches or two hemal spines. If each fused centrum is counted as two 
centra, a count of 10 + 23 = 33 vertebrae is obtained for the paratype. This count 
is in the range of the total number of vertebrae (33-35) for other specimens of 
this species. The low anal-fin ray count may be a result of the same factor that 
caused the fusions, which would make this count anomalous as well. 

Comparisons. — Greenfield (1979) proposed that Starksia ocellata be recognized 
as a superspecies containing six allospecies, including S. brasiliensis. These al- 
lospecies are distinguished with certainty only by color pattern, all meristics and 
morphometrics overlapping to some extent among the species. Based on meristics 
and head color pattern, S. brasiliensis is most similar to a species group that 
includes S. occidentalis of the western Caribbean, S. variabilis of the southwestern 
Caribbean, and S. guttata of the southeastern Caribbean and the southern islands 
of the Lesser Antilles. Within this group, S. brasiliensis has a head color pattern 
that differs from that of S. guttata only in having a narrower Y-shaped marking 
on the cheek. Starksia brasiliensis also differs from S. guttata in having a mode 
of 16 scales in the arched portion of the lateral line (vs. 17), a mode of 20 scales 
in the straight portion of the lateral line (vs. 21), and a mode of 36 total lateral 
line scales (vs. 38, all counts for S. guttata based on Greenfield 1979). The two 
species overlap in each of these counts, but the least overlap occurs in the number 
of scales in the arched portion of the lateral line. Both S. brasiliensis and S. guttata 
have a dark area on the lips, restricted to the anterior tips of the jaws, and lack 
the dark bars on the lips found in S. occidentalis and S. variabilis (S. culebrae of 
the Lesser Antilles also has bars on the lips). Starksia brasiliensis differs from all 
of these in having a narrow Y-shaped mark on the cheek. Modes for all meristic 
characters are the same for S. brasiliensis, S. occidentalis, and S. variabilis. An 
additional character that distinguishes S. brasiliensis from, at least, S. occidentalis 
and S. variabilis is the color of the ring-like marks on the cheek. Greenfield (1979) 
stated that these marks were white in S. occidentalis and lemon yellow in S. 
variabilis. In our specimens of S. brasiliensis, these marks are bright red. The 
color of these marks has not been reported for S. guttata or S. culebrae, but it is 
orange in S. ocellata from the Tortugas (Longley and Hildebrand 1941) and from 
the eastern Gulf of Mexico (pers. observ. by JTW). 

A cursory survey of the gill-raker counts for Starksia specimens in the Florida 
State Museum suggests that S. brasiliensis has fewer gill rakers on the first arch 
than other species in the S. ocellata superspecies. A specimen of S. occidentalis 
has a total gill-raker count of 10 (left)-9 (right), one S. variabilis has 9-9, and one 
S. ocellata has 11-11. This character also shows overlap as S. brasiliensis has 7 
to 9 (one specimen with 9) gill rakers, but may prove useful in distinguishing 
some of the allospecies. 

Additional support for the recognition of Starksia brasiliensis as a valid species 
in the S. ocellata superspecies comes from its geographic isolation from other 
allospecies. Collete and Riitzler (1977) suggested that the fresh silt-laden waters 
of the Amazon and Orinoco rivers might serve as a barrier to dispersal of reef 
fishes confined to depths shallower than about 50 m. Although one specimen in 
the British Museum (Natural History) was collected from a cable in 55-62 m, 


VOLUME 96, NUMBER 4 643 


there is some doubt as to the accuracy of this depth recording. This specimen was 
collected around 1890 when depths presumably were determined by taking sound- 
ings at regular intervals, thus the 55 and 62 m depths probably correspond to two 
soundings taken at separate points along the cable. Since the bottom topography 
between the two soundings 1s unknown, it is conceivable that a rocky or coralline 
outcropping on the bottom could significantly alter the 55-62 m depth range. 
Aside from this one specimen, no specimens belonging to the S. ocellata super- 
species have been reported from depths greater than 25 m. One of us (JTW) has 
collected S. ocellata at a depth of about 36 m in the eastern Gulf of Mexico 
(specimens deposited in University of South Alabama Ichthyological Collection), 
but this is still much shallower than the Brazilian record. We feel that it would 
be premature to suggest that S. brasiliensis commonly occurs at depths of 50 m 
or more until additional collections are made at these depths. In view of the 
typically shallow (less than 13 m) depth distribution of S. brasiliensis, it seems 
likely that the Amazon-Orinoco barrier effectively isolates S. brasiliensis from its 
northern congeners, but additional collecting is needed to confirm its absence 
from this region. 

In view of the distinctness of the Brazilian population from other populations 
of Starksia based on coloration, pigment pattern, and certain meristic characters, 
we agree with Greenfield (1979) and recognize S. brasiliensis as a valid species 
in the S. ocellata superspecies. 

Material examined.—CAS-SU 7750 and 53510 (holotype and paratype, re- 
spectively, of Brannerella brasiliensis) both from near Maceio, Brazil, 1899. GCRL 
9476 (2) Brazil, Bahia, Isla Itaparica, Barra do Gil, 13°00’00’S, 38°37'00’W, 
tidepools just inside outer edge of reef, rock-sand bottom, algae, bryozoa, 25 Aug 
1972, 0-2 m. BMNH 1890.1.27.19(1), Brazil, 11°50’S, 38°47'W, from cable in 
55-62 m. UF 32889(4), Brazil, Ilha das Couves, 23°25'05”S, 44°51'50’W, coral 
encrusted rocks on sand bottom, 18 Jul 1981, 10-13 m. MZUSP uncat. (3), same 
data as UF 32889. UF 32887 (1), Brazil, Ilha da Vitoria, 23°44’40’S, 45°01'40"W, 
bottom with large boulders encrusted with sponges, small corals, and sparse algal 
growth, 25 May 1981, 10-13 m. MZUSP uncat. (1), same data as UF 32887. UF 
32888 (3), Brazil, Ilha da Vitoria, 23°44’S, 45°01’W, bottom with large boulders 
encrusted with corals and sponges, 14 Jun 1981, 10-13 m. MZUSP uncat. (4), 
same data as UF 32888. 

Comparative material examined.—Starksia ocellata: UF 16018 (1), Delray 
Beach, Palm Beach County, Florida. Starksia occidentalis: UF 23348 (1 paratype), 
Frenchman’s Cay, Belize. Starksia variabilis: UF 23349 (1 paratype), Santa Marta, 
Colombia. 


Acknowledgments 

We thank William N. Eschmeyer and Tomio Iwamoto for allowing us to borrow 
the type specimens from CAS, and Charles E. Dawson for the loan of GCRL 
specimens. Alwyne Wheeler kindly provided information about the method of 
determining depth for the specimen at the BMNH. James A. McGrath and Ray- 
mond Dee, assisted in taking radiographs. We particularly thank Rosemea De 
Souza Smart, who made the Brazilian collections possible. Victor G. Springer and 
Carter R. Glbert kindly reviewed an earlier draft of the manuscript and provided 
helpful comments. Examination of the specimen at the BMNH was made possible 


644 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


by NSF grant DEB 8207313 (predoctoral dissertation improvement grant) to 
J. T. W. The typescript was prepared by Grace Russell. 


Literature Cited 


Bohlke, J. E., and V. G. Springer. 1961. A review of the Atlantic species of the clinid fish genus 
Starksia.— Proceedings of the Academy of Natural Sciences of Philadelphia 113(3):29-60. 

Collette, B. B., and K. Riitzler. 1977. Reef fishes over sponge bottoms off the mouth of the Amazon 
River. — Proceedings, Third International Coral Reef Symposium, Rosenstiel School of Marine 
and Atmospheric Science, University of Miami, May 1977:305-310. 

George, A., and V. G. Springer. 1980. Revision of the clinid fish tribe Ophiclinini, including five 
new species, and definition of the family Clinidae.—Smithsonian Contributions to Zoology 
307:1-31. 

Gilbert,C.H. 1900. Results of the Branner-Agassiz Expedition to Brazil. III. The fishes. — Proceedings 
of the Washington Academy of Sciences 2:161-184. 

Greenfield, D. W. 1979. A review of the western Atlantic Starksia ocellata-complex (Pisces: Clinidae) 
with the description of two new species and proposal of superspecies status. — Fieldiana Zoology 
73(2):9-48. 

Longley, W. H., and S. F. Hildebrand. 1941. Systematic catalogue of the fishes of Tortugas, Florida 
with observations on color, habits, and local distribution. — Carnegie Institution of Washington 
Publication 535, Papers from Tortugas Laboratory 34:1—331. 

Steindachner, F. 1876. Ichthyologische Beitrage (V).—Sitzungsberichte der Kaiserlichen Akademie 
der Wissenschaften in Wien 74 (1-2, no. 1):49—240. 


Florida State Museum and Department of Zoology, University of Florida, 
Gainesville, Florida 32611. (JTW: Present address, Division of Fishes, National 
Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560.) 


PROC. BIOL. SOC. WASH. 
96(4), 1983, pp. 645-657 


DESCRIPTION OF A NEW SPECIES OF ECHIODON 
(TELEOSTEI: CARAPIDAE) FROM ANTARCTIC AND 
ADJACENT SEAS 


Douglas F. Markle, Jeffrey T. Williams, and John E. Olney 


Abstract.— Echiodon cryomargarites, a previously unrecognized species of car- 
apid fish, is described from adult and larval specimens collected in relatively deep 
water from the Antarctic and adjacent seas. This species differs from E. exsilium 
and E. dawsoni, which have fewer precaudal vertebrae, and from EL. dentatus and 
E. drummondii which have slightly higher D,, counts, fewer pectoral rays, and 
fewer vertebrae to the origin of the dorsal fin. The new species has a relatively 
deep, flaccid body and was collected over bottom depths ranging from 400-1200 
m off Argentina, South Georgia Island, and New Zealand. Two other specimens 
that differ slightly in geographic and depth distribution, appearance, and have a 
shallow, firm body were caught at depths of 1500-1666 m off southern Chile. 

The distal tips of the transverse processes of the first two vertebrae in E. 
cryomargarites are ligamentously attached to paired sclerified structures of the 
anterior swimbladder. This condition may represent a precursor to the rocker 
bone specialization of Onuxodon. 


During independent investigations of the pearlfish genus Echiodon (Olney and 
Markle 1979; Williams and Shipp 1982), a previously undescribed species was 
encountered in samples from southern oceans. The species was apparently men- 
tioned by Andriashev (1977) and a larva now assigned to this species was collected 
by Dr. Thor Mortensen from Campbell Island shepherds (Rendahl 1926). 

In this report we examined 26 adults and five larvae of this deep-dwelling, 
southern ocean Echiodon. Our purpose is to document their distribution and 
describe the new species we recognize. Echiodon, as recognized herein, is defined 
by an eclectic group of characters taken from Arnold (1956), Cohen and Nielsen 
(1978), Robertson (1975), Maul (1976), Olney and Markle (1979) and Williams 
and Shipp (1982). For the purposes of this paper, the genus is diagnosed as those 
carapids! with 15—25 pectoral fin rays, 20—29 precaudal vertebrae, a free maxillary, 
enlarged canines at jaw symphyses, an anus posterior to a vertical through the 
pectoral fin base, no median rocker bone at anterior end of swimbladder, and 
larvae in which the first dorsal ray inserts immediately posterior to the vexillum 
and the vexillum origin is posterior to a vertical through the first anal ray. Hy- 
potheses on phylogenetic relationships within Echiodon are beyond the scope of 
this paper, but such studies are in progress. 


1 We have been informed by G. S. Myers through G. C. Steyskal that the stem of the family name 
is Carap—rather than Carapod—(Steyskal 1980) since the derivation is from a native Brazilian word, 
¢arapo, and not the Greek, pous. Thus, the family is Carapidae not Carapodidae. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


646 


Fig. 1. Echiodon cryomargarites, holotype, LACM 10985-6. A, Left lateral view; B, View of head; 
Echiodon specimen, LACM 42593-2. C, Left lateral view; D, View of head. 


Methods and Materials 


Counts and measurements of adults follow Williams and Shipp (1982), and of 
larvae follow Olney and Markle (1979). Abbreviations D3, and Aj4 refer to the 
dorsal and anal rays whose bases lie anterior to vertebra 31. 

Cephalic lateralis pore series terminology used in this paper follows Williams 
and Shipp (1982) except as follows. The lateral series (LT) is restricted to include 
those large pores of the lateral-temporal canal between that canal’s junctions with 


VOLUME 96, NUMBER 4 647 


Table 1.—Summary of selected measurements for adult E. cryomargarites and E. specimens. Values 
for holotype listed in parentheses. All measurements given as percent of head length. 


E. cryomargarites E. specimens 
Snout length 19-27 (20) 18-21 
Orbit diameter 23-29 (23) 24-25 
Iris diameter 18-24 (20) 18-19 
Upper jaw length 54-66 (56) 56-58 
Lower jaw length 55-59 (55) 54—56 
Predorsal length 160-200 (180) 180-190 
Preanal length 120-150 (130) 120-130 


the preopercular (POP) and the infraorbital (IO) canals. The supratemporal series 
(ST) includes those large pores on both sides of the head that are dorsad of the 
junctions of the supratemporal canal with the lateral temporal canals. Pores were 
located by directing a jet of air through the canals. 

Gosline (1960) pointed out that there is confusion about the definition of ophi- 
diiform “ribs,” since some of the anterior transverse processes articulate basally 
with their respective vertebral centra (‘sessile ribs”’ of Regan 1912), while pos- 
terior transverse processes fuse basally to their centra. Since pyramodontines have 
pleural ribs at the ventral ends of elongate transverse processes, we prefer to avoid 
the term “ribs.” The term transverse process used herein follows Gosline (1960) 
and is equivalent to the “sessile ribs” and transverse processes of Regan (1912) 
and the “ribs” of Courtenay and McKittrick (1970). 

Cranial vault length refers to the greatest anterior to posterior distance in a 
sagittal plane within the braincase as determined from radiographs and cleared 
and stained specimens. 

Material examined is listed under individual species accounts. Museum acro- 
nyms used are LACM—Los Angeles County Museum, Los Angeles, California; 
ISH— Institut fiir Seefischerei, Zoological Museum, University of Hamburg, Ham- 
burg, West Germany; ZMUC— Zoological Museum, University of Copenhagen, 
Copenhagen; and USNM—National Museum of Natural History, Smithsonian 
Institution, Washington, D.C. Other abbrevaitions used are TL—total length, 
SL—standard length, IK MT—Isaacs—Kidd Midwater Trawl, SO—supraorbital se- 
ries, [O—infraorbital series, LT — lateral series, ST —supratemporal series, POP— 
preopercular series, and MD—mandibular series. 


Echiodon cryomargarites, new species 
Figs. 1A and B 


Holotype.—LACM 10985-6, 410 mm TL, 37.3 mm HL, male; “‘E/tanin” sta 
1422, 56°19’—21'S, 158°29’E, 12 Feb 1965, 3 m (10 ft) Blake trawl, 833-842 m 
bottom depth range. 

Paratypes.— Off New Zealand: LACM 10985-8, 3 specimens, 230-310 mm TL, 
21.8—27.3 mm HL, males; caught with holotype. LACM 10985-7, damaged; caught 
with holotype. LACM 10979-4, ca. 220 mm TL, ca. 16 mm HL, female; “E/tanin” 
sta 1414, 52°17'—22'S, 160°40'-34’E, 9 Feb 1965, 3 m Blake trawl, 659-798 m. 


648 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


-@ -------~g 


LE as 


<= 

> 

x< 
ee 


CR 


oe 


an ae 


Fig. 2. Echiodon cryomargarites, LACM 10985-6, diagrammatic illustration of cephalic lateralis 
pores and upper lip lobes. A, left lateral view; B, Dorsal view; C, Ventral view of lower jaw; D, 
Magnified snout region showing lobes under upper lip. Prepared by W. Zomlefer. 


LACM 10984-3, three specimens, ca. 210 mm TL (female), ca. 220 mm TL 
(female), 255 mm TL (male), 16.0—22.7 mm HL; “Eltanin” sta 1419, 54°32'— 
31'S, 159°02’-O1’E, 10 Feb 1965, 3 m Blake trawl, 494-714 m. 

Off Argentina and Brazil: ISH 317/71, 280 mm TL, 20.5 mm HL, female; 


VOLUME 96, NUMBER 4 649 


0 


(0) 5 10 15 20 25mm 
eS ee |e jee 


Fig. 3. Echiodon cryomargarites, A, B. Drawings of male holotype, LACM 10985-6, from off New 
Zealand. C, D. Drawings of female paratype ISH 300/71 from off Falkland Islands. Prepared by W. 
Zomlefer. 


“Walther Herwig” sta 312/71, 46°53’'S, 60°00'W, 18 Feb 1971, bottom trawl, 800 
m. ISH 1108/66, four specimens, 181—243 mm TL, 12.3-17.0 mm HL, females; 
“Walther Herwig” sta 230/66, 35°04’S, 52°15'W, 12 Jun 1966, bottom trawl, 600 
m. ISH 1554/66, 113+ mm TL, 12.3 mm HL, male; “Walther Herwig” sta 428/ 


650 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


A Transverse Processes 1-3 


Tenth 
cranial nerve 


First vertebra 


Exoccipital 
foramen 


Fig. 4. Echiodon cryomargarites, LACM 10984-3. A, Right lateral view of first four centra illus- 
trating modified transverse processes of first three centra, position of tenth cranial nerve, and swim- 
bladder modifications; B, Ventral view of A. 


66, 34°48’S, 52°02’W, 28 Jul 1966, bottom trawl, 400 m. ISH 1229/66, 209 mm 
TL, 12.0 mm HL, female; “‘Walther Herwig’’ sta 268/66, 39°56’'S, 55°58’W, 19 
Jun 1966, bottom trawl, 600 m. ISH 1580/66, three specimens, 188-245 mm 
TL, 12.7-20.0 mm HL, females; “Walther Herwig” sta 438/66, 33°41'S, 51°12'W, 
31 Jul 1966, bottom trawl, 600 m. ISH 1818/68, two specimens, 268-302 mm 
TL, 22.3-23.3 mm HL, males; “‘Walther Herwig’ sta 65/68, 29°57'S, 47°35'W, 
27 Feb 1968, bottom trawl, 1200 m. Off South Georgia Is.: ISH 334/76, 255 mm 
TL, 19.8 mm HL, male; “Walther Herwig” sta 33/76, 53°36'S, 35°40'W, 11 Dec 
1976, bottom trawl, 800 m. 

Off Falkland Is.: ISH 300/71, two specimens, 310-315 mm TL, 25.7—27.8 mm 
HL, females; “ Walther Herwig’ sta 241/71, 55°00'S, 57°50'W, 4 Feb 1971, bottom 
trawl fished in midwater above a bottom of 775-850 m. 

Diagnosis.—A species of Echiodon from southern oceans with a relatively flac- 


VOLUME 96, NUMBER 4 651 


Table 2.—Selected measurements and counts of E. cryomargarites larvae. All measurements as 
percent head length. 


ZMUC USNM ZMUC 
Dana 3641 I 257747 LACM 42592-1 LACM 11505 P7710 
Head length 4.6 mm 4.4 mm 7.9 mm 3.6 mm 7.0 mm 
Snout length 26 23 20 — 28 
Eye diameter M2) DI DP) — 24 
Upper jaw length 44 — 51 — — 
Lower jaw length 56 55 58 64 57 
Vexillum length 236 218 311 492 — 
Snout to vexillum 196 198 233 DY 220 
Snout to dorsal 203 207 247 238 236 
Snout to anal 169 171 184 151 156 
Snout to anus 155 155 186 153 200 
Azo 45 45 48 45 — 
D3 39 40 36 34 — 
Vertebrae to vexillum 10 10 12 13 11 
Vertebrae to dorsal 11 11 13 14 12 
Vertebrae to anal 8 8 9 8 8 
Precaudal vertebrae — De — — — 


cid body, caught over depth range 400-1200 m, D;,. 36-40, two pores in POP 
canal. 

Description. — Morphometric data are summarized in Table 1. Meristic data are 
as follows, holotype in parentheses: D,, 36—40 (38), A3 9 46-50 (47), pectoral fin 
rays on left side 19-21 (21), anal fin origin under vertebrae 6—8 (6), dorsal fin 
origin over vertebrae 11-12 (11), precaudal vertebrae 25-29 (28), developed gill 
rakers 3, olfactory rosette with seven pairs of lamellae (clear and virtually un- 
countable in holotype and paratypes of LACM 10985), pores of cephalic lateralis 
system: SO-5, IO-9, LT-2, ST-3, POP-2, MD-5 (generally difficult to see), upper 
lip bears three or four small lobes (Fig. 2). Sensory papillae arranged in a single 
row along most sensory canals, and, although difficult to discern, on snout and 
top of head. 

The general appearance of E. cryomargarites is shown in Figs. 1 and 3. The 
body is flaccid, overall color in ethanol is tan with relatively uniform distribution 
of small melanophores, more concentrated along myosepta and bases of vertical 
fins; tongue ranging from pale cream colored to black; skin lining mouth and 
branchial cavity ranging from tan with scattered melanophores to uniform brown 
(dense melanophores); outer surface of stomach black; outer surface of intestine 
usually black, sometimes tan; peritoneum tan with scattered melanophores. 

Posterior portion of maxillary unsheathed (Figs. | and 3); each premaxillary 
and dentary with 1-2 large canines anteriorly; teeth on premaxillaries conical, 
depressible, and pluriserial, tapering to one row anteriorly at the base of the canines 
and to two rows of shorter teeth posteriorly; dentary teeth pluriserial with inner- 
most row fixed and outer row depressible; dentary not noticeably narrowed be- 
tween anterior canines and band of conical teeth; palatines with two irregular 
rows of blunt conical teeth; vomer with an irregular row of small, conical teeth 
laterally flanking a median row of 1-5 slightly larger conical teeth; three small 


652 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 5. Echiodon cryomargarites larva, LACM 42592-1. Dotted lines represent continuation of 
dorsal and anal fins. Stippling on body used to highlight contours and myosepta and does not represent 
pigmentation. 


tooth patches on or associated with pharyngobranchials 2, 3 and 4; fifth cerato- 
branchial with medial row of small, conical teeth. 

Lateral line difficult to follow, apparently restricted to anterior one third of 
body. 

Limited osteological observations were made on three specimens cleared and 
counter stained for cartilage and bone (ISH 1108/66, ISH 1818/68, LACM 10985- 
7 and LACM 10984-3). Otoliths (sagittae) reduced relative to other carapids, their 
lengths about 10-15% of cranial vault length. Transverse processes of first three 
vertebrae with typical carapine modifications (Regan 1912; Courtenay and 
McKittrick 1970; Olney and Markle 1979). First two processes moveable, third 
and subsequent rigid (Fig. 4A). Distal tips of first two processes ligamentously 
attached to paired sclerified structures at anterior end of swimbladder (Fig. 4A 
and B). Sclerified structures stain lightly with Alcian Blue and lie anteriad of a 
constriction in swimbladder (Fig. 4B). (This condition may represent a precursor 
of the rocker bone in Onuxcodon. If so, the rocker bone may be an ossified anterior 
chamber of a swimbladder that was derived from the main swimbladder.) Third 
process expanded, curving inward, with its posterior surface attached to the swim- 
bladder behind the constriction (Fig. 4). Swimbladder tapers slightly posteriad 
and ends under vertebrae 15-16. Large tenth cranial nerve (Vagus) passes along 
vertebral column, under first two transverse processes and above remainder. 

Etymology.—A combination of the Greek “‘kryos’”’ meaning cold and “‘mar- 
garites,” a pearl, in reference to the Antarctic distribution of this new pearlfish. 


VOLUME 96, NUMBER 4 653 


Ll 


0) 5 10 15 20 25mm 
re a 


Fig. 6. Echiodon specimen, LACM 42593-1, 30.8 mm HL. A, Left lateral view; B, Close up of 
head. Prepared by W. Zomlefer. 


Echiodon cryomargarites— Larvae 


Material Examined. —Off New Zealand: ZMUC Dana 3641!, 77 mm TL, 4.6 
mm HL, 43°40’S, 176°36’E, 8 Jan 1929, stramin net, 300 m wire out. USNM 
257747, ca. 67 mm TL, 4.4 mm HL; 43°00’S, 173°57'24”E, 7 Mar 1974, 1 m net. 
ACV S05" 65-— mm iE: 32:6" mm! Als SE lianin sta 22172 S1c29,0'S; 
160°13.0’E, 20 Jun 1968, IKMT. ZMUC P7710, 190 mm TL, 7.0 mm HL; ca. 
42°S, 174°E, 10 Dec 1914, collected by Campbell Island shepherds on shore of 
Perserverance Harbor, Antarctic Convergence, mid-Pacific: LACM 42592.1, ca. 
205 mm TL, 7.9 mm HL; “Eltanin” sta 16, 49°06-10’S, 120°13-15'W, 19 Dec 
1965, IKMT, 0-200 m. 

Comments. — Larvae were identified as belonging to this species on the basis of 
their unique combination of meristic characters (A3,, D3., precaudal vertebrae, 


654 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 7. Chart showing distribution of adults and larvae of E. cryomargarites and E. specimens. 


and fin positions), distributional data, and gross similarity to larvae of E. drum- 
mondiiand E. dentatus (Maul 1976, personal observation). Selected measurements 
and counts are shown in Table 2. 

The general appearance of the largest larva is illustrated in Fig. 5. Noteworthy 
features are the sac-like drooping gut (Rendahl 1926) to which the first anal fin 
ray 1s attached, the relatively long anal fin rays, and the absence of pigment. In 
the three smallest specimens, 3.6—4.6 mm HL, about ten internal pigment spots 
were present along the vertebral column. These commence at centra 13 or 14 and 
are spaced about 7 to 12 centra apart. These were difficult to see even in the 
smallest specimen due to occlusion by epaxial musculature but were readily ap- 
parent in the 4.4 and 4.6 mm specimens after trypsin digestion. 


Echiodon Specimens 
Figs. 1C and D 


Material Examined.—LACM 42593-1, 335 mm TL, 30.8 HL, female; ““E/tan- 
in” sta 21, 53°13’-16’S, 75°41'W, 5 Jan 1966, 1.5 m Blake trawl, 1500-1666 m. 
LACM 42593-2, 300 mm TL, 25.6 mm HL, female; same data as LACM 42593- 
ile 


Comments.— These two specimens differed slightly in geographic distribution 
and morphology from E. cryomargarites. They have a firm body, D3) 35 and were 
caught at depths of 1500-1666 m. The general appearance of these specimens 1s 
shown in Fig. 6. Coloration, dentition, and otoliths are as for E. cryomargarites. 
The lateral line is difficult to follow but apparently extends over two thirds of the 
body length. Morphometric data are summarized in Table 1. Meristic data are 
as follows: D3, 35, A;) 46-48, pectoral fin rays on left side 20-21, anal fin origin 
under vertebrae 6-8, dorsal fin origin over vertebra 12, precaudal vertebrae 27— 
29, developed gill rakers 3, olfactory rosette with seven pairs of lamellae. 


VOLUME 96, NUMBER 4 655 


1600 4-Larvae rT r 
e -Echiodon cryomargarites 
= -Echiodon specimens 


800 


600 


Average Depth of Capture (m) 


400 


200 


(0) 10 20 30 40 
Head Length (mm) 


Fig. 8. Relationship between average depth of capture and size (head length) of E. cryomargarites 
adults and larvae and E. specimens. 


These specimens described above closely resemble and may be conspecific with 
E. cryomargarites. Both forms are distinguished from E. exsilium and E. dawsoni, 
which have fewer precaudal vertebrae (21-24) and from E. dentatus and E. drum- 
mondii, which have slightly higher D3, (42—45), fewer pectoral fin rays (15-17), 
and fewer vertebrae to the dorsal fin origin (8-9). 


Distribution 


The geographic distribution of E. cryomargarites is shown in Fig. 7. The well 
established dispersal capabilities of the larvae (Olney and Markle 1979) and the 
West Wind Drift are sufficient to account for the wide geographic range of the 
species. Antimora rostrata, Halargyreus johnsonii, and Merluccius australis have 
similar distribution patterns (Iwamoto 1975; Cohen 1973; Inada 1981). The first 
two of these species co-occurred with E. cryomargarites in four of eleven (36%) 
of our adult collections (M. Stehmann, personal communication, 5 Jan 1982; 
LACM catalog records). In at least one case, M. australis, geographic variants had 
been described as different species but are now recognized as synonyms (Inada 
1981). 

There was no evidence of commensalism, and the species appear to be free 
living. 

With one exception (ISH 300/71) all adults were caught in bottom trawls fished 
on continental or insular slopes. There appears to be ontogenetic descent in E. 


656 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


cryomargarites with larger specimens being found deeper (Fig. 8). The two Echio- 
don specimens were found at about twice the depth of similar-sized E. cryomar- 
garites (Fig. 8). 


Acknowledgments 


We are very grateful to the following curators and their institutions for making 
specimens available: R. Lavenberg and J. Seigel (LACM), G. Krefft and M. Steh- 
mann (ISH), E. Bertelsen (ZMUC), Les Knapp (USNM). We thank the following 
for constructive review of this manuscript: E. Anderson, R. Crabtree and J. Carter 
(VIMS) and Drs. Bob Lavenberg and Dan Cohen (LACM). Olney and Markle 
were supported in part by NSF Grant DEB-81-17185. This is a joint contribution 
of the Huntsman Marine Laboratory, the Florida State Museum and the Virginia 
Institute of Marine Science (VIMS Contr. No. 1132). 


Literature Cited 


Andriashev, A. P. 1977. Some additions to schemes of the vertical zonation of marine bottom fauna. 
Pp. 351-360 in G. A. Llano (ed.). Adaptations within Antarctic ecosystems. — Proceedings of 
the Third SCAR Symposium on Antarctic Biology. 

Arnold, D.C. 1956. A systematic revision of the teleost family Carapidae (Percomorphi, Blennioidea) 
with descriptions of two new species.—Bulletin of the British Museum (Natural History) 
Zoology 4:245-—307. 

Cohen, D. M. 1973. The gadoid fish genus Halargyreus (Family Eretmophoridae) in the Southern 
Hemisphere.—Journal of the Royal Society of New Zealand, 3(4):629-634. 

Cohen, D. M., and J. G. Nielsen. 1978. Guide to the identification of genera of the fish order 
Ophidiiformes with a tentative classification of the order.— NOAA Technical Report NMFS 
Circular 417, 72 pp. 

Courtenay, W. R., Jr., and F. A. McKittrick. 1970. Sound-producing mechanisms in carapid fishes, 
with notes on phylogenetic implications.— Marine Biology 7:131-—137. 

Gosline, W. A. 1960. Hawaiian lava-flow fishes, part IV. Snyderidia canina Gilbert, with notes on 
the osteology of ophidioid families.— Pacific Science 14(4):373-381. 

Inada, T. 1981. Two nominal species of Merluccius from New Zealand and southern South America. — 
Japanese Journal of Ichthyology 28(1):19-30. 

Iwamoto, T. 1975. The abyssal fish Antimora rostrata (Ginther).—Comparative biochemistry and 
physiology 52B:7-11. 

Maul, G. E. 1976. The fishes taken in bottom trawls by R. V. ““Meteor” during the 1967 Seamount 
Cruises in the Northwest Atlantic.— Meteor Forschungsergebnisse, Reihe D-Biologie 22:1-69. 

Olney, J. E., and D. F. Markle. 1979. Description and occurrence of vexillifer larvae of Echiodon 
(Pisces: Carapidae) in the western North Atlantic and notes on other carapid vexillifers.— 
Bulletin of Marine Science 29(2):365-379. 

Regan, C. T. 1912. The classification of the blennioid fishes.—Annals and Magazine of Natural 
History, (8), 10:265—280. 

Rendahl, H. 1926. Papers from Dr. Th. Mortensen’s Pacific Expedition 1914-16. XXX. Fishes from 
New Zealand and Aukland-Campbell Islands.— Videnskabelige Meddelelser fra Dansk Natur- 
historisk Forening 81:1—14. 

Robertson, D. A. 1975. Planktonic stages of the teleost family Carapidae in eastern New Zealand 
waters.— New Zealand Journal of Marine and Freshwater Research 9:403—409. 

Steyskal, G. C. 1980. The grammar of family-group names as exemplified by those of fishes. — 
Proceedings of the Biological Society of Washington 93(1):168-177. 

Williams, J. T., and R. L. Shipp. 1982. A new species of the genus Echiodon (Pisces: Carapodidae) 
from the eastern Gulf of Mexico.—Copeia 1982(4):845—-851. 


(DFM) The Huntsman Marine Laboratory, St. Andrews, New Brunswick, Can- 
ada EOG 2X0; (JTW) Florida State Museum and Department of Zoology, Uni- 


VOLUME 96, NUMBER 4 657 


versity of Florida, Gainesville, Florida 32611; present address: Division of Fishes, 
NHB WG-12, Smithsonian Institution, Washington, D.C. 20560; (JEO) Virginia 
Institute of Marine Science and the College of William and Mary, Gloucester 
Point, Virginia 23062. 


PROC. BIOL. SOC. WASH. 
96(4), 1983, pp. 658-663 


TELEOSTEAN OTOLITHS FROM THE LATE 
CRETACEOUS (MAESTRICHTIAN AGE) SEVERN 
FORMATION OF MARYLAND 


Richard W. Huddleston and Kurt M. Savoie 


Abstract.—Approximately 1000 teleostean fish otoliths were recovered from 
the Severn Formation, Late Cretaceous (Early—Middle Maestrichtian) of Mary- 
land. These otoliths represent at least 14 kinds of fishes belonging to eight families 
(Pterothrissidae, Argentinidae, Ariidae, Ophidiidae, Polymixiidae, Trachichthyi- 
dae, ?Pempheridae, and Apogonidae) and three unidentified families, suborders 
Albuloidei, Stomioidei, and Anguilloidei. Otoliths of Vorhisia sp. dominated the 
fauna, representing approximately 54 percent of the total identified otoliths. The 
next dominant form, represented by the Apogonidae, comprised 27 percent of 
the identified fauna. 


Previous description of teleostean otoliths from the Late Cretaceous of North 
America are limited. Frizzell (1965a) described Prealbula weileri and Protalbula 
sohli, based on isolated sagittae from the Earliest Campanian, Eutaw Formation 
of Alabama. Frizzell (1965b) described Vorhisia vulpes, based on isolated lapilli 
from the Maestrichtian, Fox Hills Sandstone Formation of South Dakota. Frizzell 
and Koenig (1973) described asterisci from the same formation and assigned them 
to V. vulpes. These asterisci, however, do not belong to Vorhisia (J. E. Fitch, pers. 
comm.). Huddleston (1981) described Bernardichthys zorraquinosi, based on sa- 
gittae from the Early Cenomanian, Bernard Formation of Oregon. 

The present study is based on approximately 1000 otoliths recovered from 
about 100 kg of fossiliferous matrix collected from an exposure at the base of the 
Severn Formation by one of us (KMS). All field samples were collected from 
LACM (Los Angeles County Museum, section of Vertebrate Paleontology) locality 
4425; Beltway exit 34W, Central Avenue, Prince George County, Maryland. The 
locality was initially exposed by the cutting action of a small unnamed creek. 
Construction activities have increased the exposure. Samples were taken from a 
30 cm thick section of scattered shell and small lenses of dark gray shell marl, 
mixed with broken shell. 

The term Severn Formation was first proposed by Denton (1891) for a variety 
of lithologic units. Clark, Bagg and Shattuck (1897) later proposed the term Mon- 
mouth Formation for certain lithologic units occurring in New Jersey. This term 
also was applied to Late Cretaceous marine strata in Maryland. Recently Minard, 
Shol, and Owens (1978) reintroduced the term Severn Formation to replace the 
term Monmouth Formation in Maryland. The Severn Formation is Late Creta- 
ceous, Early-Middle Maestrichtian and corresponds to the Navarroan Provincial 
stage (Brouwers and Hazal 1978). 


VOLUME 96, NUMBER 4 


Fig. 1. A, B, Pterothrissidae, left sagitta, 15.4mm, LACM 4425/120101 (A-innerface; B-outerface); 
C, Pterothrissidae, right sagitta, 11.5mm, LACM 4425/116980; D, E, Anguilloidei-A, left sagitta, 
10.5mm, LACM 4425/116981, (D-innerface; E-outerface); F, G, Albuloidei, left sagitta, 7.4mm, 
LACM 4425/116979 (F-innerface; G-outerface). 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 2. A, cf. Pempheridae, left sagitta, 4.0mm, LACM 4425/1 16990; B, Polymixiidae, right sagitta, 
4.5 mm, LACM 4425/116988; C, Anguilloidei-B, right sagitta, 6.1mm. LACM 4425/116983; D, E, 
Ariidae, right lapillus, 4.9mm, LACM 4425/116985 (D-innerface; E-outerface); F, G, Vorhisia sp., 
left lapillus, 25mm, LACM 4425/116984 (F-innerface; G-outerface). 


VOLUME 96, NUMBER 4 


Fig. 3. A, Ophidiidae, right sagitta, 4.0mm, LACM 4425/116987; B, Argentinidae, left sagitta, 
3.0mm, LACM 4425/116983; C, near-Apogonidae-B, right sagitta, 2.5mm, LACM 4425/116994; D, 
Stomioidei, left sagitta, 3.1mm, LACM 4425/116993; E, Argentinidae, left sagitta, 2.79mm, LACM 
4425/120115; F, near-Apogonidae-A, left sagitta, 2.5mm, LACM 4425/116991; G, Argentinidae, left 


sagitta, 3.42mm, LACM 4425/120116; H, near-Apogonidae-A, right sagitta, 3.42mm, LACM 4425/ 
116992. 


662 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Taxa Number of otoliths Figure 
Pterothrissidae 12 fig. 1A—C 
Albuloidei 13 fig. 1F—G 
Anguilloidei-A 2 fig. 1D-E 
Anguilloidei-B 2 fig. 2C 
Argentinidae 3 fig. 3B, E, G 
Stomioidei 4 fig. 3D 
Ariidae: Vorhisia sp. 596 fig. 2F—G 
Ariidae: unidentified 82 fig. 3A 
Polymixiidae 11 fig. 2E 
Trachichthyidae 63 not figured 
c.f. Pempheridae 1 fig. 2A 
Apogonidae-A 190 fig. 3F, H 
Apogonidae-B 106 fig. 3C 

Discussion 


The ichthyofauna of the Severn Formation as defined by the otoliths, was 
dominated by the family Ariidae, with approximately 54 percent of the otoliths 
of a single genus, Vorhisia sp. Waage (1968) noted Vorhisia as a brackish and 
freshwater indicator. While Vorhisia sp. dominated the ichthyofauna all of these 
otoliths displayed varing degrees of attrition. The extent of potential postmortem 
transportation is not determinable. It is possible that these otoliths were washed 
in from a shallower area. However, it is unlikely that this Vorhisia sp. represented 
a freshwater species. 

The albuloid sagittae were uncommon in the Severn samples and while the 
Recent Albulidae are found in warm subtropical to tropical environments it cannot 
be construed with any degree of certainty that the Severn albuloids occupied 
similar habitat without additional data. Additionally all of the albuloid otoliths 
displayed degrees of erosion and these otoliths may have undergone considerable 
postmortem transport. 

The extant Polymixiidae, Apogonidae, Pempheridae, and Ariidae reflect a trop- 
ical-subtropical environment and their presence in the Severn fauna may suggest 
this condition. However, additional supportive data are necessary to confirm these 
conclusions. This late Cretaceous fauna is unusual in that its overall components 
more closely resemble otolith faunas observed from the Paleocene Brightseat 
Formation rather than the other Cretaceous age ichthyofaunas represented by 
otoliths (Huddleston, unpublished data). 


Acknowledgments 


We thank D. Haman, Chevron Oil Field Research Company for review of this 
manuscript and helpful comments, and D. Nolf, Belgian Museum of Natural 
History for his suggestions on the improvement of this study. We especially thank 
the late J. E. Fitch, San Pedro, California for review of this manuscript and for 
providing invaluable advice and comparative material throughout this study. 


Literature Cited 


Brouwers, E. M., and Hazel, J. E. 1978. Ostracoda and correlations of the Severn Formation (Na- 
varroan; Maestrichtian) of Maryland.—Journal of Paleontology (Supplement)52(2):1—52. 


VOLUME 96, NUMBER 4 663 


Clark, W. B., Bagg, R. M., and Shattuck, G. B. 1897. Upper Cretaceous deposits of Maryland.— 
Maryland Geological Survey (Upper Cretaceous):23-110. 

Denton, N. H. 1891. Mesozoic formations of Eastern Virginia and Maryland.— Geological Society 
of America, Bulletin 2:431—450. 

Frizzell, D. L. 1965a. Otolith-based genera and lineages of fossil bonefish (Clupeiformes, Albuli- 

dae). —Senckenbergiana Lethaea 46a:85-1 10. 

1965b. Otoliths of new fish (Vorhisia vulpes, n. gen., n. sp. Silurioidei ?) from the Upper 

Cretaceous of South Dakota.—Copeia 1965(2):178-181. 

, and Koenig, J. W. 1973. Upper Cretaceous Ostariophysine (Vorhisia) redescribed from 

unique association of utricular and lagenar otoliths (lapillus and asteriscus).—Copeia 1973(4): 

692-698. 

Huddleston, R. W. 1981. Bernardichthys zorraquinosi, a new genus and species of salmoniform fish 
from the late Cretaceous of Oregon. — Proceedings of the Biological Society of Washington 94(1): 
37-42. 

Minard, J. P., Sohl, N. F., and Owens, J. P. 1978. Reintroduction of the Severn formation (Upper 
Cretaceous) to replace the Monmouth formation in Maryland.—U.S. Geological Survey, Bul- 
letin 1435—A:132-133. 

Waage, K. M. 1968. The type Fox Hills formation, Cretaceous (Maestrichtian), South Dakota. Part 
1. Stratigraphy and paleonenvironments.— Bulletin of the Peabody Museum of Natural History, 
Yale University 27, 167pp. 


RWH, Scientifc Research Systems, 1 1044 McGirk, El Monte, California 91731; 
KMS, 2311 Pimmit Dr. no. 702 E, Falls Church, Virginia 22043. 


PROC. BIOL. SOC. WASH. 
96(4), 1983, pp. 664-668 


A NEW SUBSPECIES OF FOX SPARROW FROM 
ALASKA 


J. Dan Webster 


Abstract. — Passerella iliaca chilcatensis, new subspecies, is described from the 
mainland of southeastern Alaska and adjacent Canada. The action represents a 
split of the well-known race P. i. fuliginosa, from which P. 1. chilcatensis differs 
in being less reddish and duller in color and in having a shorter tail. 


Swarth (1920) in his monograph on the Fox Sparrows and emendation of the 
range of P. i. fuliginosa (1922) intimated that the race might need subdivision. 
In earlier publications (Webster 1950, 1975) I, too, lumped the population of the 
mainland of southeastern Alaska and adjacent parts of Yukon and British Co- 
lumbia with those of Vancouver Island and northwestern Washington. A similar 
arrangement was followed by Munro and Cowan (1947), the American Orni- 
thologists’ Union (1957), and Gabrielson and Lincoln (1959). In view of differ- 
ences in color, size, and migration patterns, I now conclude that a formal de- 
scription of what Swarth (1920) called “‘non-typical fuliginosa’’ is required. 


Passerella iliaca chilcatensis, new subspecies 


Holotype.—California Academy of Sciences #70787, collected 12 Jun 1981 by 
J. Dan Webster, 7 miles (airline) south-southwest of Klukwan, Alaska, near Tsirku 
River, about 250 m elevation, in alder-willow thicket near large cottonwood trees. 
Little body fat; 37.8 g, male, cloacal protuberance large, testes 13 and 10 mm 
long; adult skull. 

Subspecific characters. — Blacker, less reddish than P. i. townsendi, both dorsally 
and on ventral spots; less reddish, more sepia (or yellowish), duller than P. 7. 
fuliginosa, but equally dark or blackish both dorsally and on ventral spots, and 
with shorter tail. Dorsum of fresh-plumaged birds nearest fuscous, though slightly 
more olivaceous than that; compared directly with Smithe (1975). 

Breeding range. —Chilkat River area of British Columbia and Alaska southeast 
along the mainland to the Tewart area of British Columbia. Precise localities, 
including those of intergradient populations, are listed below. Thickets, mostly 
of alder bushes but sometimes mixed with small spruces or willows, comprise the 
breeding habitat. These are mostly near timberline, but also occur in logging 
clearings, avalanche rubble, river margins, down even to sea level. 

Winter range.—Most specimens I examined were from the coast of Oregon 
(Tillamook) and northern California south to San Francisco. However, one spec- 
imen had been taken in Alaska—Craig, Prince of Wales Island, 5 Jan 1920; and 
one on Puget Sound, Washington, 10 Jan 1912. Also, specimens intermediate 
between chilcatensis and one or another of the adjacent breeding races were 
examined. These had been taken in winter (Dec—Feb) in south coastal British 
Columbia and south to Tulare County, California. Swarth (1920) examined spec- 
imens of “‘non-typical fuliginosa” from the coastal counties of northern California 


VOLUME 96, NUMBER 4 665 


Table 1.—Passerella iliaca. Length of wing. Males, breeding populations. 


Subspecies n r M SD CV 
P. i. chilcatensis 51 76-86 80.235 2.08 2.59 
P. i. fuliginosa 14 79-85 82.357 1.99 2.41 
P. i. altivagans 19 77-85 82.053 2.19 2.67 
P. i. zaboria 50 81-92 87.220 1.65 1.89 
P. 1. annectens 12 78-86 82.583 2.26 2.74 
P. i. townsendi 70 75-86 80.814 2.38 2.94 


south to the San Francisco Bay region, plus single specimens from San Luis Obispo 
and San Bernardino counties farther south. Grinnell and Miller (1944), although 
stating that the winter range in California was concentrated in the humid north- 
west, cited additional specimens from Monterey, Los Angeles, Siskiyou, Shasta, 
and Lassen counties. It is notable that the winter range of fu/iginosa (sensu stricto) 
is confined to British Columbia and Washington. 

Measurements.—Tables 1-4 give a comparison of the new race with adjacent 
races. Wing chord, tail, and exposed culmen were measured in standard fashion, 
but depth of bill by Swarth’s method (1920:83). Only adult male specimens taken 
10 May to 8 Aug were included in the tables. In summary, chilcatensis differs 
significantly in size from adjacent races only on these points: zaboria has a longer 
wing and shorter bill; fuliginosa has a longer tail. While annectens has a deeper 
bill, the difference is not significant. Within the samples of the races chilcatensis 
and townsendi (as classified in the tables of measurements), some specimens from 
geographically marginal areas showed intermediacy of color characters. 

Discussion. —I have listed (Tables 1—4) five races as breeding adjacent to chil- 
catensis—annectens to the northwest, townsendi to the west, zaboria to the north- 
east, altivagans to the east and southeast, and fuliginosa to the south. However, 
there is certainly a possibility of contact between chilcatensis and olivacea some- 
where in the coast range of British Columbia, as suggested by a few winter spec- 
imens (cf. Phillips 1964). 

In view of the finding by Zink (1982) of slight but significant size differences 
in a Califronia deme of Passerella iliaca in samples separated by 54 years of time, 
the size differences reported herein must be viewed with caution. This caution 
applies especially to the race annectens, in which 10 of the 12 male specimens I 
examined were collected 90 years ago, whereas for the other races a high proportion 


Table 2.—Passerella iliaca. Length of tail. Males, breeding populations. 


Subspecies n Tr M SD CV 
P. i. chilcatensis 49 62-75 69.306 2.53 3.66 
P. i. fuliginosa 14 70-81 75.143 2.42 3D 
P. i. altivagans 18 68-72 73.556 4.21 5/2 
P. i. zaboria 50 67-76 71.420 2.11 2.95 
P. 1. annectens 12 67-73 70.250 1.88 2.67 


P. i. townsendi 69 65-78 71.072 DAD 3.87 


666 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Table 3.—Passerella iliaca. Length of culmen. Males, breeding populations. 


Subspecies n Tr M SD CV 
P. i. chilcatensis 51 11.0-13.5 12.249 0.54 4.45 
P. i. fuliginosa 14 11.1-13.0 12.221 0.56 4.55 
P. i. altivagans 19 11.0-12.5 11.674 0.42 3.60 
P. i. zaboria 48 10.0-12.3 11.308 0.55 4.87 
P. i. annectens 12 LOIN) 11.600 0.28 2.44 
P. i. townsendi 69 10.1-13.2 11.990 0.64 5.36 


were collected within the last 20 years. Gabrielson and Lincoln (1951) commented 
on “‘foxing”’ in Alaskan Fox Sparrow specimens; I concur that postmortem changes 
in color, mostly from black towards red, are considerable. Comparisons of spec- 
imens for color were made only with specimens of approximately the same mu- 
seum age. 

According to Gabrielson and Lincoln (1959) and earlier workers, the Fox Spar- 
row does not breed on the southern islands of the Alexander Archipeligo of 
Southeastern Alaska except for Wrangell and Forrester islands. However, I noted 
two singing birds near Ketchikan, Revillagigedo Island, 9 Jun 1977, and about 
25 in logging clearcuts on Prince of Wales Island, 9-24 Jun 1977. While I found 
no nests in the latter area, I collected nine adults (of townsendi) in breeding 
condition. (The one female had a prominent brood patch and had recently laid 
an egg.) It is probable that the species did breed on these islands prior to the onset 
of major logging operations 20 years ago, but only in small numbers and at high 
elevations near timberline. The few observers before 1959 (see reports by Bailey, 
Swarth and Willett, as cited by Gabrielson and Lincoln 1959) spent little or no 
time at high elevations during summers. On 22 Jun 1977 my assistant, Keith 
Gehring, climbed a high ridge above the logging clearcut in the valley of Beaver 
Creek, Prince of Wales Island. He saw one singing Fox Sparrow at an elevation 
of about 2000 feet. Apparently, the recent clearcut logging operations on Prince 
of Wales and Revillagigedo islands, as on the mainland near Haines and Hyder, 
have allowed a great population expansion of the Fox Sparrow. Near Haines, I 
found the race chilcatensis breeding in considerable numbers (three nests found) 
in timberline alder thickets at several locations in Jun and Jul 1972, 1975 and 
1981 (cf. Webster 1975). Banks (1970) reported expansion of range of two other 
subspecies of Passerella iliaca in Oregon after clearcut logging. 


Table 4.—Passerella iliaca. Depth of bill. Males, breeding populations. 


Subspecies n iP M SD CV 
P. i. chilcatensis 51 7.8—9.6 8.745 0.48 5.45 
P. i. fuliginosa 13 8.5—9.9 9.031 0.43 4.80 
P. i. altivagans 19 8.5-9.5 9.021 0.28 3.08 
P. i. zaboria 48 8.4-9.8 9.100 0.33 3.62 
P. i. annectens 1. 8.2-10.1 9.233 0.54 5.84 
P. i. townsendi 69 8.0-9.7 8.809 0.40 4.55 


VOLUME 96, NUMBER 4 667 


Breeding season specimens examined.—(If a locality is marked +, specimens 
from there are intermediate toward the adjacent race.) Passerella iliaca chilca- 
tensis: ALASKA 37—7 miles SSW Klukwan; 8 miles SSW Klukwan, on Porcupine 
Mountain; 4—5 miles W Klukwan, near Klehini River; Four Winds Mountain, 
above Mosquito Lake; Chilkat Mountain; Glacier Station, on White Pass railroad; 
Mountains above Juneau; 7-8 miles up (=NE) Stikine River from Point Rothsay; 
Berg Creek, 15 miles ESE Wrangell; 4 miles NW Hyder. YUKON 6—miles 97- 
98 Haines Highwayt. BRITISH COLUMBIA 38—near Bear Glacier, 15 miles 
NE of Stewart; Snowbank Pass, 47 miles NW Meziadin Junction; 24 miles S 
Kinaskan Lake, near Iskut River; miles 46-56 Haines Highway, Rainy Hollow; 
mile 85 Haines Highwayt+; Doch-da-on Creek; Flood Glacier; Great Glacier, on 
Stikine River. . 

Passerella i. annectens: ALASKA 17—Yakutat; Situk River; Osier Island. 

Passerella i. zaboria: ALASKA 33— Many localities in N and W Alaska; Lower 
Tonsina; Chistochina. YUKON 24—Several localities in N and central Yukon; 
Tagisht. BRITISH COLUMBIA 5—Dease Lake; Atlin. NORTHWEST TER- 
RITORIES 1—Niultin Lake. MANITOBA 5—Churchillf. 

Passerella i. townsendi: ALASKA 60—Glacier Bayt; Lemesurier Island; Chich- 
agof Island; Admiralty Island; St. Lazeria Island; Baranof Island; Wrangell Island?*; 
Forrester Island; Prince of Wales Island. BRITISH COLUMBIA 41 — Queen Char- 
lotte islands; Goose Island; Bella Coola. 

Passerella i. altivagans: BRITISH COLUMBIA 37—Nine-mile Mountain; 
Hudson Bay Mountain; Mount Cronin; Driftwood Mountains; Parsnip River; 
Williams Lake; above Stuie; Mount Revelstoke; Glacier National Park. 

Passerella i. fuliginosa: BRITISH COLUMBIA 16—Several localities on Van- 
couver Island; Triangle Island; Alert Bay. WASHINGTON 3—Tatoosh Island; 
Neah Bay; La Push. 


Acknowledgments 

My field work was supported by the Hanover College Faculty Research Com- 
mittee and the Indiana Academy of Science. The British Columbia Provincial 
Museum, the National Museum of Canada, the Museum of Comparative Zoology 
of Harvard University, the Royal Ontario Museum of Zoology, the San Diego 
County Museum, and the National Museum of Natural History loaned me spec- 
imens for study. The California Academy of Sciences, the Field Museum of 
Natural History, the University of Michigan Museum of Zoology and the Museum 
of Vertebrate Zoology of the University of California allowed me to study in their 
collections. 


Literature Cited 
American Ornithologists’ Union. 1957. Check-list of North American birds. 5th ed. American 


Ornithologists’ Union, Lancaster, Pa. 691 pp. 
Banks, R. C. 1970. The Fox Sparrow on the west slope of the Oregon Cascades.— Condor 72:369- 


370. 

Gabrielson, I. N., and F. C. Lincoln. 1951. Post-mortem color changes in bird specimens. — Condor 
53:298-299. 
, and . 1959. The birds of Alaska. Stackpole Company, Harrisburg, Pa. 922 pp. 


Grinnell, J., and A. H. Miller. 1944. The distribution of the birds of California.— Pacific Coast 
Avifauna 27:1-608. 


668 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Munro, J. A., and I. McT. Cowan. 1947. A review of the bird fauna of British Columbia.— British 
Columbia Provincial Museum, Special Publications 2:1—285. 

Phillips, A. 1964. Remarks on Fox Sparrows.—Jn Phillips, A., J. Marshall, and G. Monson. The 
birds of Arizona. University of Arizona Press, Tucson. 220 pp. 

Smithe, F. B. 1975. Naturalist’s color guide, part I. American Museum of Natural History, New 
York. 

Swarth, H.S. 1920. Revision of the avian genus Passerella, with special reference to the distribution 

and migration of the races in California.— University of California Publications in Zoology 

21(4):75—224. 

1922. Birds and mammals of the Stikine River region of northern British Columbia and 
southeastern Alaska.— University of California Publication in Zoology 24(2):125-314. 
Webster, J. D. 1950. Notes on the birds of Wrangell and vicinity, southeastern Alaska.— Condor 
52:32-38. 

1975. The Fox Sparrow in southwestern Yukon and adjacent areas.—Condor 77:215-216. 
Zink, R.M. 1983. Evolutionary and systematic significance of temporal variation in the Fox Sparrow. — 

Systematic Zoology 32:223-238. 


Hanover College, Hanover, Indiana 47243. 


PROC. BIOL. SOC. WASH. 
96(4), 1983, pp. 669-674 


A REVISION OF THE GOLFINGIA SUBGENERA 
GOLFINGIELLA, STEPHEN, 1964, AND 
STPHONOIDES, MURINA, 1967 
(SIPUNCULA) 


Edward B. Cutler, Norma J. Cutler, and Peter E. Gibbs 


Abstract.—The Golfingia subgenera Golfingiella and Siphonoides are regarded 
as void since all of the included species have either been transferred to other taxa 
(G. immunita, G. pudica, and G. mexicana) or placed on the list of species in- 
quirenda (G. approximata and G. innoxia) or incertae sedis (G. quadrata). 


Cutler and Murina (1977) began the process of reviewing the sipunculan genus 
Golfingia and its included subgenera: subsequent work has led to an examination 
of the difficult Apionsoma/Mitosiphon/Fisherana complex (Cutler 1979) and Thy- 
sanocardia (Gibbs, Cutler & Cutler, in press). The present paper continues this 
survey with an assessment of the Golfingia subgenera Golfingiella Stephen, 1964, 
and Siphonoides Murina, 1967. Experience has shown that many published de- 
scriptions contain serious errors and for this reason, every attempt has been made 
to examine type-materials. Taxa are here regarded as analogous to scientific hy- 
potheses; if they cannot, for whatever reason, be tested (verified or falsified), then 
they should no longer be considered as valid entities. 

The literature references under each species name are only those in which 
material is newly recorded. For additional references see Stephen and Edmonds 
(1972). Table 1 lists the currently accepted names of the species considered here 
with their proposed status. Four of the six species were erected on single specimens. 


Genus Golfingia Lankester, 1885 
Subgenus Golfingiella Stephen, 1964 


Golfingiella species are characterized as having two pairs of introvert rectractor 
muscles, introverts without hooks, single-lobed nephridia and a posteriorly-at- 
tached spindle muscle. 

The subgenus was originally established by Stephen (1964) for five little-known 
species of Golfingia which Fisher (1950) had omitted from his revision. Phas- 
colosoma approximatum is named as the type-species but Stephen makes no 
mention of the other species included. In Stephen and Edmonds (1972) four species 
are listed: G. pusilla (Sluiter), G. abnormis (Sluiter), G. innoxia (Sluiter), and G. 
approximata (Roule); G. trichocephala was transferred from Golfingiella to Gol- 
fingia sensu stricto. 

Cutler and Murina (1977) reviewed Golfingia and made a few changes in Gol- 
fingiella: G. abnormis was synonymized under Phascolosoma pectinatum Kefer- 
stein. Two species (G. pudica (Selenka) and G. immunita (Sluiter)) were moved 
into this subgenus after examining type-material and the literature. Golfingia 
pusilla was synonymized under G. trichocephala placing them in the subgenus 


670 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Table 1.—The proposed status of the species assigned to Golfingiella and Siphonoides. Species with 
an asterisk are based on single specimens. 


Currently accepted name Proposed status 
*Golfingia (Golfingiella) approximata (Roule, 1898) species inquirenda 
*Golfingia (Golfingiella) immunita (Sluiter, 1902) Golfingia (Apionsoma) 
immunita 
*Golfingia (Golfingiella) innoxia (Sluiter, 1912) species inquirenda 
Golfingia (Golfingiella) pudica (Selenka, 1885) 
sensu Selenka, Stephen Golfingia (Golfingia) 
cf. margaritacea 
sensu Wesenberg-Lund, Murina, Cutler Golfingia (Apionsoma) 
(cf. immunita) 
Golfingia (Siphonoides) mexicana Murina, 1967 Aspidosiphon (Aspidosiphon) 
(includes A. longirhyncus) mexicanus 
*Golfingia (Siphonoides) quadrata (Ikeda, 1905) incertae sedis 


Mitosiphon. Subsequently Mitosiphon was submerged under Apionsoma (Cutler 
1979). These actions leave the four species listed in Table | to consider. 


Golfingia (Golfingiella) approximata (Roule, 1898) 


Phascolosoma approximatum Roule, 1898:385; 1906:77-81, pl. 9, fig. 87, pl. 10, 
figs. 100-101. 


Type-locality. —Off Moroccan coast at 1105 m depth. 

The single representative of this species in the Paris museum (Cat. #V23) is 
not in good condition. The internal organs are somewhat disrupted and since 
most of the introvert is missing, nothing can be added regarding the hooks, spines, 
or tentacles. There are four retractor muscles. Whether or not the spindle muscle 
is attached to the posterior end of the trunk cannot be determined: a short piece 
of white thread-like tissue is at the posterior end which might be the remnant of 
a spindle muscle or possibly it is a fragment of the ventral nerve cord. Subsequent 
collections of benthic organisms in the area of the type locality have yielded no 
additional specimens of this species. Due to the damaged, incomplete state of the 
holotype and only specimen, the accuracy of Roule’s description cannot be verified 
and therefore, G. approximata is placed on the list of species inquirenda. 


Golfingia (Golfingiella) immunita (Sluiter, 1902) 
Phascolosoma immunitum Sluiter, 1902:40-41. 


Type-locality.—Indonesia, 6°N, 121°E, 275 m. 

This single 8 mm worm with a 6 mm introvert (not 3 and 8 mm as reported) 
is in the Amsterdam museum (Cat. #V. Si. 198). As Sluiter reported, it does have 
a posteriorly-attached spindle muscle and lacks longitudinal muscle bands. How- 
ever, introvert hooks in rings, each bearing small basal spinelets (Fig. 1) are present 
and there are four introvert retractor muscles (Sluiter states that it has no hooks 
and only two retractors). The general impression one gets from its external form 


VOLUME 96, NUMBER 4 671 


Fig. 1. Introvert hook from the holotype of Golfingia (Apionsoma) immunita. Scale line equals 10 
micrometers. 


is that of a Phascolosoma or a papillated Golfingia (Apionsoma). As the introvert 
is partially withdrawn, the tentacles are difficult to discern but they appear to be 
arranged around the nuchal organ, as typical for Golfingia (Apionsoma). 

This individual clearly belongs in the subgenus Apionsoma and 1s similar to G. 
capitata in overall form but differs in hook structure. Therefore, the species name 
is referred to this subgenus. 


Golfingia (Golfingiella) innoxia (Sluiter, 1912) 
Phascolosoma innoxium Sluiter, 1912:13, text-fig. 3a—b. 


Type-locality.— Azores, 35°25'N, 31°22’W, 1229 m. 

As noted in Cutler and Murina (1977) the single specimen on which Sluiter 
based this species is housed at the Musée Océanographique in Monaco but is not 
available for close examination. There are no drawings of its internal anatomy 
and it is difficult to accept Sluiter’s statements without being able to verify them. 
Many collections have been made in the general vicinity of the type locality over 
the past 70 years but no additional worms matching this description have been 
reported. 

In view of doubts concerning the validity of this species (individual) the name 
is here on the list of species inquirenda until further information can be obtained 
to adequately determine its status. 


Golfingia (Golfingiella) pudica (Selenka, 1885) 


Phascolosoma pudicum Selenka, 1885:11-—12, figs. 14-16.— Fischer, 1929:484.— 
Stephen, 1948:217-218. 


672 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Golfingia pudica.— Wesenberg-Lund, 1959:59-60, text-fig. 4.—Murina, 1976:65.— 
Cutler and Cutler, 1979:954—955. 


Type-locality.— Kerguelen Islands, 18-225 m. 

This enigmatic species has been recorded five times by five different authors. 
These reports and specimens (in chronological order) are as follows: 

1885 —Selenka’s original description made no mention of the posterior attach- 
ment of the spindle muscle. Examination of the type-material (from Kerguelen) 
in the British Museum (Nat. Hist.) showed that this muscle is not attached to the 
posterior end of the trunk and, therefore, belongs in Go/fingia sensu stricto. When 
Cutler and Murina (1977) moved it to Golfingiella they erroneously assumed 
Wesenberg-Lund’s (1959) interpretation to be correct. Selenka compared G. pud- 
ica to G. vulgaris but maintained its distinctiveness based on hooks being in rows, 
not scattered, papillae distribution (on introvert as well as trunk), and number of 
tentacles (20 or less). The eight worms in the type collection are all less than 18 
mm (trunk) and not all have hooks; these look very much like G. margaritacea. 
Smaller individuals of G. margaritacea from Japan do have deciduous hooks 
(Cutler, Cutler & Nishikawa, in preparation). Fischer’s (1929) account is merely 
a repeat of Selenka’s material with no new information. 

1948—Stephen recorded over 100 specimens from Kerguelen but this material 
cannot be located in any British museum. Stephen comments, “In many cases, 
the body wall was in rather degenerate condition, so that the distinguishing char- 
acters could be made out only with difficulty.” There are no figures nor additional 
morphological comments. Due to the poor quality of the material it is difficult 
to accept this record. 

1959— Wesenberg-Lund reported a single 8 mm worm from Mauritius which 
has been examined in Copenhagen. It does have the posteriorly-attached spindle 
muscle as she stated and therefore cannot be G. pudica. It closely resembles 
Golfingia (Apionsoma) species in having hooks with small, basal spinelets. 

1976—Murina reported a single 16 mm worm from the East China Sea with a 
posteriorly attached spindle muscle. As noted above, this indicates it cannot be 
G. pudica and must therefore be Golfingia (Apionsoma) species. Murina states 
that the hooks are in rings but makes no mention of basal spinelets. 

1979—When Cutler & Cutler reported their six small (2-12 mm) specimens 
from the Mozambique Channel they noted that they were not in good condition 
and “‘Therefore these specimens are assigned to this taxon with reservations.” 
Similarities of G. pudica to Golfingia capitata were pointed out, and that a clear 
view of the tentacular arrangements is necessary to differentiate with certainty 
between the two. The hooks of these specimens do have rudimentary spinelets 
as shown for G. immunita (Fig. 1). 

We now propose that the three most recent collections be considered as a 
Golfingia (Apionsoma) species (possibly G. immunita). Those from Kerguelen 
should be considered as Golfingia sensu stricto and perhaps Golfingia margari- 
tacea or G. ohlini, both common in the Antarctic. 


Subgenus Siphonoides Murina, 1967 


Murina (1967) established the new subgenus Siphonoides for four species (G. 
immaniata [sic], G. mexicana Murina, G. quadrata (Ikeda), and G. rutilofusca 


VOLUME 96, NUMBER 4 673 


(Fischer)), differing from other subgenera in having only one pair of introvert 
retractor muscles and a posteriorly attached spindle muscle. The only other taxa 
with this combination of characters are found in the family Aspidosiphonidae. 

Stephen and Edmonds (1972) placed G. rutilofusca in the subgenus Phascoloides 
(now Nephasoma) because it was found that the spindle muscle is unattached. 
Golfingia immunita was moved to Golfingiella (Cutler and Murina 1977) because 
the type-specimen had four retractor muscles, not two (see above). The following 
two species thus remain to be considered. 


Golfingia (Siphonoides) mexicana Murina, 1967 


Golfingia mexicana Murina, 1967:1333-1334, fig. 3. 
Aspidosiphon longirhyncus Cutler and Cutler, 1980:4—6, figs. 4-5. 


Type-locality.— Gulf of Mexico. 19°N, 76°W, 110 m. 

A close comparison of specimens kindly provided by Dr. V. V. Murina pointed 
to the striking similarity of G. mexicana to Aspidosiphon longirhynchus Cutler 
and Cutler (1980). Unfortunately no specimen of either population is preserved 
with tentacular crowns expanded. The diameter of the introvert is very small and 
attempts to dissect out the tentacular crown met with limited success; a few 
tentacles arranged dorsal to the mouth can be discerned in 4. /ongirhyncus but 
for G. mexicana no conclusion could be reached by the authors nor by Murina. 

While acknowledging that the shields are poorly developed it is concluded that 
these two taxa are nevertheless conspecific and belong in the genus Aspidosiphon. 
Therefore, Golfingia mexicana is hereby transferred to Aspidosiphon and now 
includes 4. /ongirhyncus as a junior synonym. 


Golfingia (Siphonoides) quadrata (Ikeda, 1905) 
Phascolosoma quadratum Ikeda, 1905:170-171, pl. 8, fig. 14. 


Type-locality.—South Negros, Philippines. 

The single specimen Ikeda used as the basis for this species could not be located 
in Japan. Since some of the descriptions of Ikeda have been found to be in error, 
this species remains suspect. Cutler and Cutler (1981) pointed to its close similarity 
to G. mexicana. Specifically nothing is known about the tentacular array and the 
precise anus/nephridia location cannot be ascertained. The epidermal structures 
need to be reexamined but cannot be. No additional specimens have been assigned 
to this taxon. Ikeda (1905) compared his species to Phascolosoma macer which 
has been shown to belong to Aspidosiphon (Cutler and Murina, 1977:183). In 
view of the above, the taxonomic position of this species cannot be determined 
so this name is added to the list of incertae sedis. It is suspected that Ikeda’s 
worm was an atypical Aspidosiphon, a common genus in the Philippines. 


Acknowledgments 


This work was made possible by the kind cooperation of the following persons: 
J. Renaud-Mornant, Muséum National d’Histoire Naturelle, Paris; S. van der 
Spoel, Inst. voor Taxonomische Zool., Amsterdam; G. Testa, Inst. Oceanographie, 
Monaco; R. Sims, Brit. Mus. (Nat. Hist.), London; J. Kirkegaard, Zool. Mus., 


674 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Copenhagen; and G. Murina, Inst. of Biol. Southern Seas, Sevastopol. Funding 
has been provided by U.S. National Science Foundation grant #DEB-8011121. 


Literature Cited 


Cutler, E. B. 1979. A reconsideration of the Gol/fingia subgenera Fisherana Stephen, Mitosiphon 
Fisher, and Apionsoma Sluiter (Sipuncula).— Zoological Journal of the Linnean Society 65:367— 
384. 

Cutler, E. B., and N. J. Cutler. 1979. Madagascar and Indian Ocean Sipuncula. — Bulletin du Muséum 
National D’Histoire Naturelle, Paris, 4° ser. 1 (sec A., #4);941—990. 


, and . 1980. Sipuncula from Southern Brazil.—Boletim do Institut Oceanografico, Sao 
Paulo 29(1):1-8. 
, and 1981. A reconsideration of Sipuncula named by I. Ikeda and H. Sato.—Pub- 


lications of the Seto Marine Biological Laboratory 26:51-93. 

Cutler, E. B., N. J. Cutler, and T. Nishikawa. [in prep.] Systematics and distribution of Japanese 
Sipuncula. 

Cutler, E. B., and V. V. Murina. 1977. On the sipunculan genus Golfingia Lankester, 1885.— 
Zoological Journal of the Linnean Society 60:173-189. 

Fischer, W. 1929. Die Sipunculiden, Priapuliden und Echiuriden der Arktis.— Jn Romer, F., and F. 
Schaudinn, Fauna Arctica 5 (II):451—490. 

Fisher, W. K. 1950. The sipunculid genus Phascolosoma.— Annals and Magazine of Natural History 
(12) 3:547-552. 

Gibbs, P. E., E. B. Cutler, and N. J. Cutler. [in press] A review of the sipunculan genus Thysanocardia 
Fisher. — Zoologica Scripta. 

Ikeda, I. 1905. Gephyreans collected by Professor Dean at Manjuyodi, Southern Negros (Philippine 
Islands).— Annotations Zoologicae Japonenses 5:169-174. 

Murina, V. V. 1967. Report on the sipunculid worms from the sublittoral zone of Cuba and the 
Mexican Gulf. —Zoologichesku Zhurnal 54(9):1329-1339. 

—. 1976. New and rare species of sipunculids from the East China Sea.— Vestnik Zoologii 2: 
62-67. 

Roule, L. 1898. Note préliminaire sur les espéces de gephyriens recueillies dans les explorations sous 
marines du ‘Talisman’ et du “Travailleur’.— Bulletin du Muséum National D’Histoire Naturelle, 
Paris 4:384-387. 

Selenka, E. 1885. Report on the Gephyrea collected by H.M.S. ‘Challenger’ during the years 1873-— 
1876.—Report of Scientific Research of the Challenger 13(36):1—25. 

Sluiter, G. P. 1902. Die Sipunculiden und Echiuriden der Siboga-Expedition, nebst Zusammenstel- 
lung der Uberdies aus den indischen Archipel bekannten Arten.—Siboga Expeditie, Monogra- 
phie 25:1-53. 
—. 1912. Gephyriens (Sipunculides et Echiurides) provenant des campagnes de la Princesse 
Alice. 1898-1910.—Resulats des Campagnes Scientifiques de Prince Albert I 36:1-36. 
Stephen, A.C. 1948. Sipunculids.—Report of B.A.N.Z. Antarctic Research Expedition 1929-1931, 
(5B)4:213-—220. 

—. 1964. A revision of the classification of the phylum Sipuncula.—Annals and Magazine of 
Natural History (13)7:457—462.. 

Stephen, A. C.,and S.J. Edmonds. 1972. The phyla Sipuncula and Echiura. London: British Museum 
(Natural History) 528 pp. 

Wesenberg-Lund, E. 1959. Sipunculoidea and Echiuroidea from Mauritius.— Videnskabelige Med- 
delelser fra Danske Naturhistorisk Forening i Kjobenhavn 121:53-73. 


(EBC, NJC) Utica College of Syracuse University, Utica, New York 13502, 
U.S.A.; (PEG) Marine Biological Association U.K., The Laboratory, Citadel Hill, 
Plymouth, PL1 2PB, United Kingdom. 


PROC. BIOL. SOC. WASH. 
96(4), 1983, pp. 675-683 


TWO SPECIES OF TYLOS AUDOUIN FROM CHILE, 
WITH NOTES ON SPECIES OF TYLOS 
WITH THREE FLAGELLAR ARTICLES 
(ISOPODA: ONISCOIDEA: TYLIDAE) 


George A. Schultz 


Abstract.—Two species of Tylos Audouin are described from Chile including a 
new species, 7. chilensis, based on specimens formerly called T. spinulosus Dana. 
New specimens which more nearly match the description of 7. spinulosus are 
redescribed under that name. The two species, both with three flagellar articles 
on the flagellum of antenna 2, are compared with each other and with the three 
other species of Ty/os which also have three flagellar articles. 


Roman (1977:110) listed 24 species of Ty/os Audouin (1826) (see Ferrara and 
Taiti, 1979:91, for explanation of use of Tylos Audouin, 1826), three of which 
have three flagellar articles on antenna 2. To Roman’s list can be added T. 
marcuzzil Soika, and T. wegeneri Vandel (see Schultz 1974) making a total of at 
least 26 species in the genus (possible synonyms among them are not considered 
here). 7y/los wegeneri has three flagellar articles and 7. marcuzzii has four. The 
new species described herein and 7. wegeneri bring to five the total number of 
known species of 7y/os with three flagellar articles on the flagellum of antenna 2. 
With length and location they are: T. spinulosus Dana (1853), to 27 mm, central 
and southern Chile; 7. neozelanicus Chilton (1901), to 14 mm, New Zealand; T. 
exiguus Stebbing (1910), to 4.5 mm, Red Sea; 7. wegeneri Vandel (1952), to 22 
mm, West Indies and west coast of Costa Rica; and 7. chilensis, new species, to 
12 mm, central Chile. 

Only slight morphological differences separate most species of 7y/os (except T. 
wegeneri which has a unique pleotelson) and the species with three flagellar articles 
on antenna 2 (except 7. wegeneri) are no exceptions. 

The new species described here is based on the specimens called 7. spinulosus 
(Dana) by Schultz (1970). Specimens of a different species (also with three flagellar 
articles) kindly sent to me by Dr. Eduardo Jaramillo, Universidad Austral de 
Chile, proved to be more nearly like 7. spinulosus Dana than the specimens 
described by Schultz under that name, and they are described here as 7. spinulosus. 
I thank Dr. Eduardo Jaramillo for sending me the specimens so that accurate 
identifications could be made of the two species of 7y/os from Chile. I also thank 
Jersey City State College for separately Budgeted Research funds. 


Tylos spinulosus Dana, 1853 
Figs. 1A-J, 2A—F, 3A—-E 


Tylus spinulosus Dana, 1853:717, pl. 47, fig. la—c. 

Tylos spinulosus Dana.— Miers 1877:675.—Budde-Lund 1879:9; 1885:279; 1908: 
78.—Stebbing 1893:424.—Chilton 1901:121; 1910:288.—Van Name 1924:192; 
1936:415, 416, fig. 257. 

nec Tylos spinulosus (Dana).—Schultz 1970:302, figs. 18-27. 


676 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 1. Tylos spinulosus, male 20 mm long: A, Lateral view; B, Frontal view; C, Antenna 1; D, 
Underside pleon; E, Posterior View, F, Right mandible; G, Left mandible; H, Maxilla 1; I, Antenna 
2: J, Endopod male pleopod 2. 


Specimens have not been recorded since Dana’s record. Van Name (1936:415, 
Fig. 257) quoted Dana’s (1853) description in full and reproduced all of his 
illustrations. The specimens described here have a flagellum with the lengths of 
the articles more nearly matching those of the specimen illustrated by Dana (cf. 
Fig. 11 here with Pl. 47, Fig. 1b of Dana, also in Van Name 1936:415, Figa2am): 


VOLUME 96, NUMBER 4 677 


Fig. 2. Tylos spinulosus: A, Maxilliped; B, Maxilla 2 with detail of exopod; C, Hypopharynx; 
D-F, Male peraeopods I, II and VI. 


They also lack the notch in the posterolateral corner of peraeonal segment I as 
shown by Schultz (1970). Dana did not illustrate or record such a notch. It probably 
would have been noted by Dana if it had been present as it is large and not 
characteristic of the other species of 7y/os known at the time. 7y/os spinulosus 
sensu Schultz (1970) is herein considered a new species. 

Description. —Eyes large with many ocelli. Body with anterior peraeonal seg- 


678 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 3. Tylos spinulosus: A, Male peraeopod VII; B—E, Female peraeopods I, II, VI and VII. 


ments with low tubercules; dorsum covered with minute hairlike scales. Frontal 
plate of cephalon with length of lateral margins less than half width of plate; dorsal 
margin obtusely rounded. Antenna | pear-shaped with many aesthetascs near tip. 
Antenna 2 short; flagellum of 3 articles, longer than last peduncular segment. 
Article 1 of flagellum about twice as long as articles 2 and 3 combined; article 3 
tiny. Clypeus rounded. Mandibles each with 3 cusps on incisor process. Lacinia 
mobilis on right mandible with 3 teeth and no compound setae. Lacinia mobilis 
on left mandible with many small apical teeth arranged in crown; small setae 
present. Five setae in setal row on right mandible. Molar process of each mandible 
broad and fringed with many tiny setae. Exopod of maxilla 1 with many large 
and small teeth and one compound seta present on tip; endopod with 3 long 
medially pointing setae and small seta on tip. Hypopharynx bilobed. 

Peraeonal segment I with broadly rounded posterolateral margin. Edge of pe- 


VOLUME 96, NUMBER 4 679 


Fig. 4. Tylos chilensis: A, Lateral view anterior part; B, Frontal process; C, Antenna 2; D, Antenna 
1; E, Right mandible; F, Left mandible; G, Maxilla 2; H, Underside of pleon; I, Posterior end. 


raeonal segment II broadly rounded. Edges of peraeonal segments III—-VII some- 
what squarish. Peraeopods of male and female with very similar pattern of setae 
on inner margins. Posterior peraeopods of male (see VI and VII of male here, 
Figs. 2F and 3A) with very broad, short setae when compared to those on re- 
spective peraeopods of female (see VI and VII of female, Figs. 3D and 3E). 

Pleon with edges of all segments reaching to general body margin. Pleotelson 
projecting slightly beyond general body margin, with rounded posterior border 
and shallow lateral notches receiving edges of pleonal segment 5. Inner edges of 
right and left pleonal segment 5 extensions pointed and almost touching medially; 
inner edges of pleonal extensions of segment 4 pointed and wide apart. 

Measurements.— Males to 27 mm long; females to 26 mm long. 

Etymology.—The name spinulosus—‘‘with spinules,’—refers to the minute 
spines on the dorsum. 

Type-locality.—Nassau Bay, Fuegia (Terra del Fuego), Chile. The species lives 
on beaches where it burrows into the sand. 

Distribution.— Playa Punta Choros (29°53’S, 71°19’W), north of Coquimbo, 


680 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Chile, south to Terra del Fuego. The specimens mentioned by Van Name (1936: 
416) (AMNH 7120) from Toto, near Valparaiso are definitely of this species and 
not as recorded by Schultz (1970). 

Disposition of specimens. —Specimens have been deposited in the United States 
National Museum—USNM 204417. Others have been retained by Dr. Jaramillo 
at the Universidad Austral de Chile. 

Affinities. — The species is unique among the members of the genus with three 
flagellar articles in that the posterolateral corner of peraeonal segment I is broadly 
rounded (see below under T. chilensis, on how it differs from that species). Ap- 
parently the two species are separated ecologically. Ty/os spinulosus is present on 
the beach where it burrows, and the new species described below lives high up 
on the shore in the spray zone. 


Tylos chilensis, new species 
Figs. 4A-I, 5A-H 


Tylos spinulosus non Dana, Schultz 1970:302, figs. 18-27. 


Description. —Eyes large with many ommatidia. Body smooth; dorsum covered 
with minute hairlike scales. Frontal plate of cephalon with 5 sides, height laterally 
about half width, with obtusely pointed dorsal side. Antenna | pear-shaped with 
many aesthetascs near tip. Antenna 2 short; flagellum of 3 articles about as long 
as peduncular segment. Flagellar article 1 about twice as long as 2 and very tiny 
3 combined. Inner margins of long peduncular segments with grooves along lengths. 
Clypeus rounded. Mandibles each with 3 sharp cusps on incisor process. Lacinia 
mobilis on right mandible with 2 sharp teeth and compound setae. Lacinia mobilis 
on left mandible with crown of small teeth on tip; compound setae present. Setal 
row with 2 setae and broad flattened molar process with many tiny setae on right 
and left mandibles. Exopod of maxilla 1 with 4 large and several small teeth on 
tip; endopod with 4 long medially pointing setae on tip. Hypopharynx bilobed. 

Peraeonal segment I (lateral view) with deeply notched posterolateral margin. 
Posterior margin (lateral view) obtusely pointed just above posterolateral notch. 
Lateral margins of peraeonal segments II-IV with squarish borders. Peraeonal 
segments V—VII with rounded lateral borders. All peraeopods of male and female 
similar, with similar patterns of many long setae on inner margin of each. 

Pleon with edges of all pleonal segments reaching general body margin. Pleo- 
telson projecting slightly beyond general body margin; posterior margin produced 
medially with deep grooves laterally placed to receive pleonal segments 5. Inner 
edges of right and left medially projecting pleonal segments 3, 4 and 5 rounded 
and wide apart medially. 

Measurements.—Specimens to 12 mm long. 

Etymology.—The name chilensis refers to the country in which the species was 
collected. 

Type-locality.—Montemar, Chile (a few kilometers north of Valparaiso). Under 
rocks and in crevices high on a cliff facing the sea in the spray zone. 

Distribution.— Known only from the type-locality. 

Disposition of types.—Type-specimens have been deposited in the National 
Museum of Natural History (Smithsonian Institution): holotype male USNM 


VOLUME 96, NUMBER 4 681 


Fig. 5. Tylos chilensis: A-D, Male peraeopods, I, II, VI and VII; E-H, Female peraeopods I, II, 
VI and VII. 


204144; allotype female USNM 204415; paratypes male and females USNM 
204416. 

Affinities.—The new species differs from all other members of the genus with 
three flagellar articles in that is the only one with a notch on the posterolateral 
edge of peraeonal segment I. It differs from 7. spinulosus, as redescribed here, in 
that there is less of a protrusion or process on the posterior margin of the pleo- 


682 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


telson. There are also differences in the shapes of, and the space between the tips 
of the internal ventral extensions of pleonal segments 3, 4 and 5. 


Other Species of 7ylos with Three Flagellar Articles 


Tylos sp. De Borre (Three flagellar articles?) The status of this species from 
Peru remains unknown (Van Name 1936:416; Schultz 1970:302). 

Tylos neozelanicus Chilton (1901:120, pl. 13, fig. 2). The species from Lyall 
Bay, New Zealand, was well described and illustrated by Chilton, but it has not 
been recorded since (Hurley 1961). The relative length of the flagellar articles on 
antenna 2 (they are each about the same length) and the regularly rounded (not 
produced) posterior margin on peraeonal segment I (lateral view) sets the species 
apart from TJ. spinulosus and T. chilensis. 

Tylos exiguus Stebbing (1910:228, pl. 23). The species, based on tiny specimens 
from the Red Sea, was adequately illustrated by Stebbing who stated that it might 
be a young individual. So far 7. exiguus is the only species of the genus with three 
flagellar articles recorded from or near the Red Sea. The three flagellar articles 
are each about the same length. The posterior margin of peraeonal segment I is 
smooth and the posterolateral angle is obtuse. 

Tylos wegeneri Vandel (1952:74, figs. 4-10). Vandel described the species in 
some detail on specimens from Isla Margarita, Venezuela. It was further recorded 
by Schultz (1974) who extended its range to Tobago and St. Martins in the West 
Indies, and compared it to other species in the genus from the New World. Several 
specimens collected on 21 September, 1957, from Puntarenas, Costa Rica, were 
identified by the author from the collections of arthropods of the Department of 
Agriculture, Florida State Museum, Gainesville, Florida. The location is on the 
west or Pacific coast of Costa Rica so the range of the species is extended from 
Caribbean to Pacific shores. The unique configuration of the pleotelson sets the 
species apart from all others in the genus. 


Literature Cited 


Audouin, J.-V. 1826. Description de l’Egypte, ou recueil des observations, et des recherches qui ont 
été fetes on Egypte pendant l’expédition de l’armée francaise. Explication sommaire des planches 
de crustacés de l’Egypte et de la Syrie. Histoire naturelle, J.-C. Savigny, Paris 1(4):77-98. 

Budde-Lund, G. 1879. Prospectus generum specierumque Crustaceorum Isopodum terrestrium. 
Copenhagen pp. 1-10. 

—. 1885. Crustacea Isopoda terrestria per familias et genera et species descripta. Hauniae pp. 
1-319. 

—. 1908. Die Landisopoden der deutschen Siidpolar-Expedition 1901-1903, mit Diagnosen 
verwandter Arten.— Deutsche Siidpolar-Expedition 9(Zool. 1):69-92. 

Chilton, C. 1901. The terrestrial isopods of New Zealand.— Transactions of the Linnean Society of 

London (2) 8(4):99-152. 

. 1910. Additions to the terrestrial Isopoda of New Zealand. — Transactions of the New Zealand 

Institute 42:286-291. 

Dana, J.D. 1853. Report on the Crustacea of the United States Exploring Expedition. Part II. United 
States Exploring Expedition during the years 1838, 1839, 1840, 1841, 1842, under the command 
of Charles Wilkes, U.S.N. 14. Crustacea (Isopoda pp. 696-805). Sherman, Philadelphia. 

Ferrara, F.,and S. Taiti. 1979. A check-list of terrestrial isopods from Africa (South of the Sahara). — 
Monitore Zoologico Italiano (N.S.) Supplemento 12(10):89-215. 

Hurley, D.E. 1961. A checklist and key to the Crustacea Isopoda of New Zealand and the subantarctic 
islands.— Transactions of the Royal Society of New Zealand 1(20):259-292. 


VOLUME 96, NUMBER 4 683 


Miers, E. J. 1877. Ona collection of Crustacea, Decapoda and Isopoda, chiefly from South America, 
with descriptions of new genera and species.— Proceeding of the Zoological Society of London 
1877:653-679. 

Roman, M.-L. 1977. Les Oniscoides halophiles de Madagascar (Isopoda, Oniscoidea).— Beaufortia 
26(334):107-152. 

Schultz, G. A. 1970. A review of species of Tylos Latreille from the New World (Isopoda, Onis- 

coidea).—Crustaceana 19(2):297-305. 

. 1974. Terrestrial isopod crustaceans (Oniscoidea) mainly from the West Indies and adjacent 

regions. I. 7y/os and Ligia.—Studies on the fauna of Cura¢ao and other Caribbean Islands 45: 

162-173. 

Stebbing, T. R. R. 1893. A history of Crustacea: Recent Malacostraca. International Science Series 

74, London pp. xv1i—466. 

. 1910. Isopoda from the Indian Ocean and British East Africa.— Transactions of the Linnean 

Society of London (2) 14:83-122. 

Vandel, A. 1952. Etude des isopodes terrestres récoltes au Vénézuela par le Dr. G. Marcuzzi, suivie 
de considérations sur le peuplement deu Continent de Gondwana. — Memoire del Museo Civico 
di Storia Naturale di Verona 3:59-203. 

Van Name, W.G. 1924. Isopods from the Williams Galapagos Expedition.— Zoologica, New York 

5(18):181-210. 

1936. The American land and fresh-water isopod Crustacea.— Bulletin of the American 

Museum of Natural History 71:1—535. 


15 Smith St., Hampton, New Jersey 08827. 


PROC. BIOL. SOC. WASH. 
96(4), 1983, pp. 684-685 


BONADUCECYTHERIDAE McKENZIE, 1977: 
A SUBJECTIVE SYNONYM OF PSAMMOCYTHERIDAE 
KLIE, 1938 (OSTRACODA: PODOCOPIDA: 
CYTHERACEA) 


K. G. McKenzie 


Abstract.—Comparison of material described from the Galapagos with that 
described earlier from Malta indicates that Bonaducecytheridae McKenzie 1977, 
is a junior synonym of Psammocytheridae Klie 1938. 


Recently, Gottwald (1980) described Psammocythere hartmanni from Santa 
Cruz in the Galapagos Islands. He noted that while confident of the family in 
which the species belonged (Psammocytheridae Klie 1938) he was hesitant about 
the generic determination but did not intend to establish a new genus because he 
had only one specimen (a mature male). 

Earlier, McKenzie (1977) had described the new family Bonaducecytheridae, 
type-species Bonaducecythere hartmanni McKenzie, 1977, on the basis of material 
collected from the island of Malta, in the Mediterranean. 

Comparing the Maltese and Galapagos taxa it is clear that they have many 
characters in common. In both, the shells are small and flattened ventrally with 
an anterior gape; the line of concrescence is irregular; muscle scars include 3 
adductors plus | frontal scar; antennules are 6-segmented, with the 4th segment 
carrying 4 dorsomedial and 6 dorso-distal bristles, while the 5th segment is bare; 
the antennae have 3 terminal endopodial claws and 3-jointed flagella; the man- 
dibles have epipods with only 1—2 Strahl(en); maxillae epipods have 2 downward- 
pointing setae; the male P1’s have curved club-like clasping ungues; P2s and P3s 
are slender with 4-segmented endopods. However, there are some differences in 
the respective hemipenes, in the P1 chaetotaxy and Gottwald (1980) did not record 
whether the seminal vesicles of his specimen were coiled within the duplicature 
as is the case in Bonaduceythere. Reference to more Galapagos material is needed 
before it can be placed in Bonaducecythere. If this were done, then Psammocythere 
hartmanni Gottwald, 1980, would become a junior homonym of Bonaducecythere 
hartmanni McKenzie, 1977. 

On shell characters, Bonaducecythere is easily separated from Psammocythere 
which is broadly rounded both anteriorly and posteriorly and has a cigar-like 
shape (Klie 1938:214, figs 733, 734). However, it does share several characters 
of the soft anatomy, notably, 6-segmented antennules with identical chaetotaxy; 
3-segmented antennal exopods; mandible epipods with a reduced (1-2) number 
of Strahl(en); maxillae with reduced epipods, bearing 2—3 downward pointing 
setae; strongly dimorphic Pls, with the terminal ungues being curved and club- 
shaped in males; and slender P2s and P3s with 4-segmented endopods. No other 
cytheracean family has this complex of characters (Hartmann and Puri 1974). 
Therefore, Bonaducecytheridae should be considered a junior synonym of Psam- 
mocytheridae. 


VOLUME 96, NUMBER 4 685 


Psammocytheridae are now held to include at least P. remanei Klie, 1936; P. 
hartmanni Gottwald, 1980; and B. hartmanni McKenzie, 1977 —from Helgoland, 
Galapagos and the Mediterranean respectively. There are two Mediterranean 
records for Bonaducecythere—that by McKenzie (1977) from Malta; and an earlier 
record of n. gen., n. sp. from the island of Naxos, Greece, which, although not 
described, was sufficiently well illustrated (Barbeito-Gonzales 1971, Plate 43b) to 
identify it with B. hartmanni. 

The preferred habitats for these three small ostracode species are all interstitial. 


Literature Cited 


Barbeito-Gonzalez, P. J. 1971. Die Ostracoden des Kiistenbereiches von Naxos (Griechenland) und 
ihre Lebensbereiche. — Mittelungen Hamburger Zoologisches Museum und Institut 67:255-326. 

Gottwald, J. 1980. Systematik, postembryonale Entwicklung, Evolution und Zoogeographie inter- 
stitieller Ostracoda (Crustacea) aus dem Pazifik. Dissertation zur Erlangung des Doktorgrades 
der Mathatmatisch-Naturwissenschaftlichen Fakultat der Georg-August-Universitat zu G6ttin- 
gen, 260 pp. 

Hartmann, G., and Puri, H. S. 1974. Summary of Neontological and Paleontological Classification 
of Ostracoda.— Mitteilungen Hamburger Zoologisches Museum und Institut 70:7-73. 

Klie, W. 1938. Krebstiere oder Crustacea III: Ostracoda, Muschelkrebse. — Die Tierwelt Deutschlands 
34:1-230. 

McKenzie, K. G. 1977. Bonaducecytheridae, a new family of cytheracean Ostracoda, and its phy- 
logenetic significance. — Proceedings of the Biological Society of Washington 90:263-273. 


Riverina College of Advanced Education, P.O. Box 588, Wagga Wagga, New 
South Wales 2650, Australia. 


Note: Recently, Gottwald (1983) renamed 7. hartmanni Gottwald, 1980, as P. santacruzensis Gott- 
wald, 1983. For the reasons given above, this species becomes Bonaducecythere santacruzensis (Gott- 
wald 1983). 


Gottwald, J. 1983. Interstitielle Fauna von Galpagos XXX. Podocopida | (Ostracoda). — Mikrofauna 
des Meeresdodens 90:621-805. 


PROC. BIOL. SOC. WASH. 
96(4), 1983, pp. 686-692 


HETEROPYRAMIS ALCALA AND 
THALASSOPHYES FERRARI, NEW SPECIES 
OF CLAUSOPHYIDAE 
(CALYCOPHORAE: SIPHONOPHORAE) 
FROM THE SOUTH PACIFIC 


Angeles Alvarino and Kenneth R. Frankwick 


Abstract.—Two new species, Heteropyramis alcala, and Thalassophyes ferrarii 
are described and compared with the other species of their genera, Heteropyramis 
maculata Moser, 1925, and Thalassophyes crystallina Moser, 1925. The material 
was obtained in the plankton collections made in the South Pacific during the 
U.S. Antarctic Research Program. 


The family Clausophyidae Totton and Bargmann, 1965, is characterized by the 
presence of the somatocyst in both anterior and posterior necotophores. This 
family includes 5 genera: Clausophyes, with two species, C. ovata (Kefferstein and 
Ehlers, 1860), C. galeata Lens and van Riemsdijk, 1908; Chuniphyes, with two 
species, C. multidentata Lens and van Riemsdijk, 1908; C. moserae Totton, 1954; 
Crystallophyes (C. amigdalina Moser, 1925); Heteropyramis (H. maculata Moser, 
1925); and Thalassophyes (T. crystallina Moser, 1925). 

The two new Clausophyidae described herein belong to the genera Heteropyr- 
amis and Thalassophyes. 


Heteropyramis alcala, new species 
Figs. 1-3 


Material.—From cruises of R/V Eltanin in 1965 and 1966 (see Table 1). Ho- 
lotype: Polygastric form (nectophore), and eudoxid form (bract + gonophore) 
from Eltanin cruise 25, sta 1697, off Chile, USNM 61064. Paratypes: 1 necto- 
phore, | bract, 2 gonophores, from E/tanin cruise 16, sta 895, S of New Zealand, 
USNM 61065. 

Etymology. — Named in honor of San Diego de Alcala, patron of the University 
of San Diego, and the sister city in Spain of San Diego, California. 

Description. — Polygastric phase (Fig. 1): The anterior nectophore forms a high 
pyramid, with a height slightly more than twice its width. It is about 5 mm high 
and 2.2 mm wide, with 5 ridges. The dorsal and 2 lateral ridges reach the apex 
of the nectophore, but the 2 ventral ridges only reach a point close to the apex. 

The hydroecium is not as deep as in H. maculata. It extends up to midlength 
of the nectophore and is closed by the 2 incomplete ventral ridges which are wider 
at the inferior part. These ridges have crests of roundish contour at the inferior 
part. 

A series of white triangular spots appear on the lateral ridges and the apex of 
the nectophore. The ventral ridges and the dorsal ridge have no opaque white 
spots. In H. maculata there are 9 of these spots, one at the apex, another at the 


VOLUME 96, NUMBER 4 687 


A 


Boge ; . 
Sun oe SEO 

B F q "a 

2 i Zi "3 


Fig. 1. Heteropyramis alcala, nectophore 5 mm high: A, Ventral view; B, Dorsal view. 


marginal edge of the lateral ridges, and 2—4 intermediate along those ridges (Totton 
1954; Totton and Bargmann 1965). 

In Heteropyramis alcala the right lateral ridge has 6 opaque white spots, one 
at the marginal edge, another at midlength of the ridge, 3 about equally spaced 
between the other 2, and another at midlength of the upper half of the ridge. The 
left lateral ridge has 7 opaque white spots, one at the marginal inferior corner, 
another at about midlength of the ridge, and 4 about equally spaced between 
those 2, plus one at the midpoint of the upper half of the ridge. Another opaque 
white spot appears at the apex of the nectophore, bringing the total number of 
Opaque white spots to 14. 

The nectosac occupies the center of the pyramid, is less than half the width of 
the nectophore and reaches higher than the midlength of the nectophore. The 
ostium, opening of the nectosac, is equal in size to the diameter of the nectosac, 
and the space from the wall of the nectophore to the opening of the nectosac is 


Table 1.—Records of Heteropyramis alcala in the South Pacific region. 


Eltanin Gear/ 


cruise Station tow Location Depth Date Local time 
16 895 MPS! 49°17'S, 162°00'E 500-250 m 7 Feb 1965 1110-1124 
25 1697 BPS? 39°56’S, 85°54'W 1000-S00m 30 Sep 1966 0732-0753 
26 1794 BPS 41°58’S, 160°06'E 1000-500 m 7 Dec 1966 1425-1449 
26 1803 BPS 47°37'S, 161°49’E 1000-500 m_ 13 Dec 1966 1052-1112 


' Multiple plankton sampler. 
2 Bathypelagic plankton sampler. 


688 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 2. Heteropyramis alcala, bract 3 mm high: A, Dorsal view; B, Left view. 


a wide velum. In H. maculata the nectosac has a height less than half but more 
than '4 of the nectophore’s length. 

Eudoxid phase: The bract (Fig. 2) is pyramid-shaped, with 4 triangular sides. 
It is about 3 mm in height, and 3 mm wide at the base. The ridges of the bract 
present thin laminar extensions. 

White opaque spots in the bract of Heteropyramis alcala are distributed as 
follows: one at the vertex of the bract, one at the base of the ridge of the pyramid, 
and one at the midlength of each of the 2 ridges which form the dorsal side of 
the bract. This distribution of the white spots differs from that of Heteropyramis 


Fig. 3. Heteropyramis alcala, gonophore 2.5 mm high: A, Lateral view; B, Dorsal view. 


VOLUME 96, NUMBER 4 689 


maculata, which displays one apical white spot and one spot at the base of each 
dorsolateral ridge. 

The phylocyst of Heteropyramis alcalais in form ofa little bottle, quite different 
from the shape of that of H. maculata. 

The hydroecial canals are shorter than those of H. maculata. 

The gonophore (Fig. 3) is pentagonal-prismatic in shape, has 5 ridges, and the 
top forms a short pyramid. One ridge has opaque white spots at the inferior 
margin and the upper margin, and the other has 3 equidistant spots along the 
ridge, from the inferior to the upper margins. Another ridge has one spot at the 
inferior margin and at midlength of the ridge, and the fourth ridge has one spot 
at the upper corner and one at the inferior part, at about 4 of the distance to the 
inferior margin. The fifth ridge has no opaque white spots. 

The closely related species H. maculata Moser, 1925 is found mainly in the 
tropical oceanic regions (Alvarino 1971), and was obtained by Leloup and Hent- 
schel (1938) west of South Georgia Islands. 


Thalassophyes ferrarii, new species 
Figs. 4-5 


Material.—From cruises of R/V Eltanin in 1966 (see Table 2). Holotype: 1 
superior nectophore and 1 inferior nectophore from E/tanin cruise 25, sta 1710, 
off Chile, USNM 61066. Paratype: 1 superior nectophore and 4 inferior necto- 
phores from E/tanin cruise 26, sta 1794, Tasman Sea, USNM 61067. 


Table 2.—Records of Thalassophyes ferrarii in the South Pacific region. 


Eltanin Gear/ 

cruise Station tow Location Depth of haul Date Local time 
25 1710 BPS 42°11'S, 86°03’W 2000-1000 m 5 Oct 1966 2020-2051 
26 1794 BPS 41°58'S, 160°06’E 1000-500 m 7 Dec 1966 1425-1449 
26 1803 BPS 47°37'S, 161°49'E 1000-500m _ 13 Dec 1966 1052-1112 


Etymology.—Named for Dr. Frank Ferrari in appreciation of his dedication 
and encouragement in behalf of plankton studies and his valuable assistance in 
developing this research. 

Description. —Polygastric phase: The superior nectophore (Fig. 4) is about 7 
mm high and 2 mm wide, or less than half as wide as its height. It has 5 ridges, 
all reaching to the apex of the nectophore, and each displaying crests. The crest 
of the dorsal ridge is the narrowest, and the crests of the ventral ridges the widest; 
widest at about the posterior third of the nectophore, forming a roundish contour 
of serrate edges. The lateral ridges have wide crests at the low part of the ridge, 
close to the ostium, ending at that region in a round edge. 

The hydroecium is shallow, reaching less than half the height of the nectophore. 

The long narrow nectosac reaches near the top of the nectophore. The space at 
the lowest part of the nectophore, between the wall of the nectophore and the 
opening of the nectosac is closed by the velum, which is wide. 

In Thalassophyes crystallina Moser, 1925, the hydroecium 1s deep and extends 
up to more than half the height of the nectophore. The ridges with crests are quite 
different in 7. crystallina and T. ferrarii. In the former, the crests are narrow or 


690 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


SE 
asa aS he 


Gua, 


Fig. 4. Thalassophyes ferrarii, superior nectophore 7 mm long: A, Lateral view; B, Ventral view. 


absent; in the latter they are wide and roundish at the lowest part, presenting 
serrate edges along the crests of the ventral ridges. The nectosac in 7. crystallina 


reaches up to half the height of the nectophore, whereas in T. ferrarii it nearly 
reaches the apex of the nectophore. 


——_—— == 
Ia Fem, 


pee 


Sa 


ae 


Fig. 5. Thalassophyes ferrarii, inferior nectophore 6 mm long: A, Lateral view; B, Ventral view. 


VOLUME 96, NUMBER 4 691 


The inferior nectophore (Fig. 5) is prismatic with an apical triangular prolon- 
gation at the dorsal side. The dorsal ridge has no crest. The lateral ridges develop 
a wing-like crest at the upper part, diminishing toward the low region, to end with 
no crest at the ostium level. The ventral ridges are pleated at the top, like two 
doors or flaps, forming the hydroecial folds, and covering the hydroecial tunnel- 
like cavity; at the low part, the ridges with crests are united by a round flap 
extending down from the region of the ostium. 

Eudoxid phase: None observed. 

The closely related species 7. crystallina is found mainly in the Antarctic and 
adjacent regions (Alvarino 1971). 

The scarcity and erratic distribution of most of the species of Siphonophorae, 
Medusae and Chondrophorae is not strange to scholars working on these groups. 
This peculiar phenomenon, pointed out by Alvarino (1971, 1981), Biggs (1977), 
Biggs, Bidigare and Smith (1981), and Sears (1953) is due to the particular behavior 
of these coelenterates. The resulting scattered aggregations of populations, the 
speed and swimming characteristics enable these animals to avoid capture by 
plankton nets. 


Acknowledgments 


We would like to express our appreciation to Drs. Izadore Barret, John R. 
Hunter, and Reuben Lasker for reading the manuscript. We are grateful to Dr. 
Frank Ferrari and Dr. Betty Landrum for their valuable assistance in providing 
additional collections. Special thanks are due to Dr. Thomas E. Bowman and Ms. 
Martha Brown for their careful editing of the paper. 


Literature Cited 


Alvarino, A. 1971. Siphonophores of the Pacific, with a review of the world distribution. — Bulletin 
Scripps Institution of Oceanography, University of California San Diego, La Jolla, California 
16:1—432. 

. 1981. Siphonophorae. Atlas del Zooplancton del Atlantico Sud-occidental. — Publicacion del 

Instituto Nacional de Investigacion y Desarrollo Pesquero [INIDEP]. Ministerio de Comercio 

e Intereses Maritimos. Subsecretaria de Intereses Maritimos. Argentina, pp. 383-441. 

Biggs, D.C. 1977. Field studies of fishing, feeding and digestion in siphonophores. — Marine Behavior 

and Physiology 4(4):261-274. 

, R. R. Bidigare, and D. E. Smith. 1981. Population density of gelatinous macro-zooplankton: 

In situ estimation in oceanic surface waters.— Biological Oceanography 1(2):157-175. 

Kefferstein, W., and E. Ehlers. 1860. Auszug aus den Beobachtungen tiber die Siphonophoren von 
Neapel und Messina angestellt in Winter 1859-60.—Nachrichten von der Gessellschaft, G6- 
tingen (23):254-262. 

Leloup, E., and E. Hentschel. 1938. Die Verbreitung der Calycophoren Siphonophoren im Siid- 
atlantischen Ozean. — Wissenschaft Ergebnissen der deutschen Atlantischen Expedition auf dem 
Forschung und Vermessunsschiff “Meteor,” 1925-1927, Biologische Sonderuntersuchungen 
12(2):1-31. 

Lens, A. D., and T. van Riemsdijk. 1908. The Siphonophora of the SIBOGA Expedition. —Siboga- 
Expeditie, monographie 38:1—130. 

Moser, F. 1925. Die Siphonophoren der Deutschen Siid-Polar Expedition 1903-1904. Zugleich eine 
neue Darstellung der ontogenetischen und phylogenetischen Entwicklung der Calycophoren und 
Physophoren. — Deutschen Siid-Polar Expedition 17, zool. 9:1-541. 

Sears, M. 1953. Notes on Siphonophores. 2. A Revision of the Abylinae.— Bulletin of the Museum 
of Comparative Zoology at Harvard College 109(1):1-119. 


692 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Totton, A.K. 1954. Siphonophora of the Indian Ocean together with systematic and biological notes 
on related species from other oceans.— Discovery Reports 27:1-161. 

, and H. E. Bargmann. 1965. A synopsis of the Siphonophora.—British Museum of Natural 
History, pp. 1-250. 


(AA) National Marine Fisheries Service, NOAA, Southwest Fisheries Center, 
P.O. Box 271, La Jolla, California 92038; (KRF) University of San Diego, Alcala 
Park, San Diego, California 92110. 


PROC. BIOL. SOC. WASH. 
96(4), 1983, pp. 693-697 


CAECIDOTEA FILICISPELUNCAE, A NEW 
TROGLOBITIC ASELLID ISOPOD FROM OHIO 


Thomas E. Bowman, III, and H. H. Hobbs, III 


Abstract. — Caecidotea filicispeluncae is described from Fern Cave, Adams Co., 
Ohio. The endopod of the male pleopod 2 lacks terminal processes except the 
cannula. Affinities with other Caecidotea species are obscure. 


Records of troglobitic asellids from Ohio are rare. Fleming (1972:252) listed 
Asellus alabamensis from Cedar Fork Cave, Adams Co. This is very probably a 
misidentification; as pointed out by Lewis and Bowman (1981:55), the identity 
of A. alabamensis Stafford (1911) is uncertain and its resolution awaits the col- 
lection of topotypes. A second record is that of Caecidotea stygia Packard from 
three intermittent streams near Cincinnati, Hamilton Co. (Bowman and Beckett 
1978). We report herein the occurrence of a new troglobitic species of Caecidotea 
from Adams Co., southern Ohio. 


Caecidotea filicispeluncae, new species 
Figs. 1-2 


Material.—Ohio, Adams Co., Fern Cave (38°42'23"N, 83°22'06”W). 12 Jun 
1980, leg. M. Flynn and H. H. Hobbs, III: 7.5 mm holotype male (USNM 195368); 
4 paratypes (USNM 195369), 4.7 mm male, ? mm male (pleon and telson missing), 
7.1 mm female (small oostegites), 4.5 mm female (small oostegites). 11 Sep 1982, 
leg. H. H. Hobbs, III: 3 paratypes (USNM 195370), 6.2 mm female (no oostegites), 
5.3 mm female (small oostegites), 4.4 mm female (small oostegites). 

Etymology. — From the Latin “‘filix, -icis’’ (fern) plus “‘spelunca” (cave), referring 
to the type-locality, Fern Cave. 

Diagnosis.— Blind, unpigmented. Body narrow, elongate, sides subparallel. An- 
tenna 1 esthete formula 4-0-1. Pereopod 1 proximal palmar process a robust 
articulated spine; mesial and distal processes unicuspate, narrowly separated, 
distal process smaller. Pereopod 4 sexually dimorphic. Male pleopod | larger than 
pleopod 2, with short apical setae; lateral margin concave with short distal setae 
and long proximal setae. Male pleopod 2 endopod tip with subapical cannula, 
but no other processes. Pleopod 3 exopod apex truncate. Pleopod 4 exopod with 
single false suture (pattern B of Lewis and Bowman 1981). 

Description. — Length of largest specimen, male holotype, 7.5 mm. Sides of body 
nearly straight; pereonites successively wider, greatest body width 1.7 mm at 
pereonite 7. Head slightly more than '3 wider than long; anterior margin shallowly 
concave; postmandibular lobes moderately developed. Pereonite | subequal in 
length to pereonites 6 and 7, about '3 longer than the subequal pereonites 2-5; 
coxa visible dorsally on pereonites 5—7. Telson 4 longer than wide in female, 2 
longer in male; sides subparallel; posterior margin broadly angular in male, with 
low but distinct caudomedial lobe in female. 

Antenna | reaching slightly beyond proximal margin of 4th segment of antenna 


694 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 1. Caecidotea filicispeluncae: A, 4.7 mm male; B, D-J, 7.5 mm male holotype; C, 6.2 mm 
female; K, 7.1 mm female. A, Head, dorsal; B, C, Telson and uropods, dorsal; D, Antenna 1; E, 
Antenna 2; F’, G, Right and left mandibles; H, Maxilla 1; 7, Maxilliped, with posterior view of endite; 
J, K, Pereopod 1. 


VOLUME 96, NUMBER 4 695 


Fig. 2. Caecidotea filicispeluncae: A, H, 6.2 mm female; B-G, I—K, 7.5 mm male holotype. A, B, 
Pereopod 4; C, Pereopod 6; D, Pleopod 1; E, Pleopod 2, anterior; F, G, Endopod of pleopod 2, oblique 
and posterior views; H, Pleopod 2; J, Pleopod 3; J, Pleopod 4; K, Pleopod 5. 


696 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Sinkhole 
Entrance 
oO 


| 4 Meters 


==) FERN CAVE 


ts Adams County, Ohio 


Fig. 3. Fern Cave, vertical profile, showing the 4 levels and the sites (stars) where Caecidotea 
filicispeluncae was collected. 


2 peduncle; flagellum with up to 10 segments, esthete formula 4-0-1. Antenna 2 
0.8 xX as long as body (6 mm in holotype); flagellum with up to 41 segments (in 
holotype). 

Mandibles with 4-cuspate incisors and lacinia; spine-row with 9 and 12 spines 
in left and right mandibles. Maxilla 1 with 5 and 13 apical spines on inner and 
outer lobes, as in other species of Caecidotea. Maxilliped with 5 retinacula on 
right and left members, endite densely setose. 

Male pereopod | propus about 0.6 as wide as long; palm defined by robust 
articulated spine representing proximal process; mesial process oblique, triangular, 
its pointed apex slightly distal to midlength of palm; separated by narrow 
U-shaped cleft from shorter distal process. Female pereopod 1 propus smaller, 
mesial and distal processes absent, palm armed with dense fringe of setules; dactyl 
with row of oblique teeth on flexor margin. 

Pereopod 4 sexually dimorphic, modified in male for clasping female during 
amplexus. Femur narrow and with smaller muscles, presumably since function 
of limb in ambulation is reduced. Carpus and propus shorter and wider to ac- 
commodate larger muscles used in clasping female; spines on flexor margins 
reduced for more efficient clasping. 

Male pleopod | protopod with 3 retinacula; endopod about *4 as wide as long, 
medial margin convex, apical margin slightly convex, armed with short naked 
setae, lateral margin concave for half its length in central part, armed with naked 
setae, setae on distal part short, those on proximal part long. 

Male pleopod 2, protopod with 1 medial seta, exopod proximal segment with 
3 lateral setae, distal segment oval, '4 longer than wide, with 9 marginal setae; 
endopod with rounded lateral basal apophysis, tip with moderately long, narrow 
cannula arising subapically medial to fissure, distal third of medial margin with 
raised striae. Female pleopod 2 with about 10 plumose marginal setae and a 
curved spine at proximal third of medial margin. Pleopod 3 with about 8 setae 
on truncate distal margin. Pleopod 4 exopod pattern B, without proximal spines. 

Uropods of male holotype nearly '4 longer than telson; peduncle nearly '4 longer 
than exopod; rami linear, exopod about twice as long as endopod, latter narrower 
than exopod. 


VOLUME 96, NUMBER 4 697 


Relationships.— The combination of three characters sets C. filicispleluncae apart 
from all known species of Caecidotea. (1) Male pleopod 1 with short apical setae 
and concave lateral margin with long setae proximally. At least nine species share 
this character. (2) Male pleopod 2 endopod tip with cannula, but no other pro- 
cesses. None of the above nine species share this character. (3) Pleopod 4 with 
single false suture. Three of the nine species agree; pleopod 4 is undescribed for 
four of the nine species. 

Further discussion of relationships at this time would be fruitless, since so many 
species of Caecidotea are inadequately known. 


Habitat 


Fern Cave is located in SE Adams Co. (38°42'23’N, 83°22'06”W). The cave is 
small (total horizontal length 18 m), yet it consists of four levels, making it 
vertically one of the most complex caves in Ohio. The passages are developed in 
dolomite (Peebles Formation of the Niagaran Series). 

The entrance to Fern Cave opens from the bottom of a prominent sinkhole 
east of Blue Creek Road between Southdown Fork and Copperas Rock Hollow, 
at an elevation of 230 m. The cave name is derived from the dense growth of 
Christmas ferns (Polystichum acrostichoides) within the sinkhole. There is no 
stream in the cave, but it is a “wet” cave because of the considerable drip input 
and the presence of small rimstone pools. The location of the entrance at the 
bottom of a sinkhole suggests that the cave receives water draining into the sink. 
There is no significant development of speleothems in Fern Cave. 

Caecidotea filicispeluncae occurs in the lowest (fourth level) of Fern Cave. Water 
trickles down the southeastern wall of the terminal room, resulting in a flowstone 
coating and several small mini-rimstone pools. Caecidotea filicispeluncae occurs 
in these pools and also on the floor of the room among gravel and allochthonous 
debris (see stars in Fig. 3). 

Some physicochemical data were taken from cave water on 16 April 1963. The 
water had been diluted by heavy runoff from recent rains, but the following results 
were obtained: Temperature 10.5°C; O, 10.9 mg/I; Specific conductance 142 wmhos/ 
cm; pH 7.4; Iron 0.78 mg/l; PO,-P 0.17 mg/l; NO,3-N 2.4 mg/l; Methyl orange 
alkalinity 2.4 meq; Total hardness 110 mg/l CaCO, . 


Literature Cited 


Bowman, Thomas E., and David C. Beckett. 1978. A redescription of the troglobitic isopod, Cae- 
cidotea stygia, from the environs of Cincinnati, Ohio (Crustacea: Isopoda: Asellidae).—Pro- 
ceedings of the Biological Society of Washington 91(1):294—302. 

Fleming, Laurence E. 1972. The evolution of the eastern North American isopods of the genus 
Asellus (Crustacea: Asellidae). Part I.—International Journal of Speleology 4:221-256. 

Lewis, Julian J., and Thomas E. Bowman. 1981. The subterranean asellids (Caecidotea) of Illinois 
(Crustacea: Isopoda: Asellidae).—Smithsonian Contributions to Zoology 335: 1-66. 

Stafford, Blanche E. 1911. A new subterranean freshwater isopod.— Pomona Journal of Entomology 
3(3):572—-575. 


(TEB) Department of Invertebrate Zoology (Crustacea), NHB 163, Smithsonian 
Institution, Washington, DC 20560; (HHH) Department of Biology, Wittenberg 
University, Springfield, Ohio 45501. 


PROC. BIOL. SOC. WASH. 
96(4), 1983, pp. 698-714 


PYCNOGONIDA OF THE WESTERN PACIFIC ISLANDS 
Il. GUAM AND THE PALAU ISLANDS 


C. Allan Child 


Abstract.— Nine species of Pycnogonida are reported from the western Pacific 
islands of Guam, Saipan, and the Palau Islands. These are: Ammothella tippula, 
new species, and Anoplodactylus chamorrus, new species, both from Guam; Ache- 
lia nana (Loman) from Saipan; and Achelia sp., Ammothella elegantula Stock, 
Tanystylum rehderi Child, Anoplodactylus pycnosoma (Helfer), Anoplodactylus 
sp., Callipallene novaezealandiae (Thomson), Seguapallene micronesica, new 
species, and Austrodecus palauense, new species, from the Palau Islands. No 
particular distributional patterns can be discerned because of the scarcity of rec- 
ords from islands of the western Pacific. 


This is the second report in a series on the virtually unknown pycnogonids of 
the Western Pacific islands. In it are described two new species, Ammothella 
tippula and Anoplodactylus chamorrus, from Guam in the Marianas, and six 
species, two of which are new; Seguapallene micronesica, and Austrodecus pa- 
lauense, plus at least three species identified only to genus for lack of appropriate 
material, from the Palau Islands in the western Carolines. A single record from 
Saipan Island in the Marianas is included here to complete the records for all 
specimens known to me. 

There are no pycnogonid records to my knowledge from the Marianas and only 
two records from the Caroline Islands, both from Stock’s (1968:10, 49) report on 
the collections made by the Galathea and the Anton Bruun. One records Hilton’s 
Ammothella pacifica from Ifaluk Atoll, and the other is a record of Anoplodactylus 
digitatus (B6hm) from Koror, Palau Islands. Neither species was found among 
the specimens reported herein. Little microfaunal sampling has been done in 
Micronesia, Polynesia, or Melanesia, and it is safe to say that many new species 
of the small littoral and shallow-water genera remain to be discovered among 
these hundreds of islands and atolls. 

Few distributional patterns are worthy of note, mainly due to the scarcity of 
collecting records for pycnogonids throughout the islands of the entire Pacific 
basin. The majority of known species reported herein have been collected at other 
Pacific island localities or at least from several Indo-Pacific localities. Achelia 
nana (Loman) has been recorded in Indonesia in addition to the Philippines and 
Japan. Ammothella elegantula Stock is known from the Straits of Malacca and 
Tanystylum rehderi was recorded from the Society Islands. Anoplodactylus pyc- 
nosoma (Helfer) has been recorded from Japan to Madagascar and from a number 
of intermediate localities, while Callipallene novaezealandiae (Thomson) has re- 
cently been noted at Enewetak Atoll, Marshall Islands (Child 1982:277), and was 
known previously from New Zealand, Australia, and off East Africa. 

The new species have their closest affinities with related species of the Pacific 
basin, as would be expected. Ammothella tippula is closest to A. pacifica Hilton, 


VOLUME 96, NUMBER 4 699 


noted above from Ifaluk, Caroline Islands, along with other Indo-Pacific distri- 
bution. Anoplodactylus chamorrus is closest to two other compact species; A. 
derjugini Losina-Losinsky, from the Sea of Japan, and A. compactus Hilton, a 
species from the California coast and islands. Seguapallene micronesica has no 
close distributional relations as the genus is known only from one other species 
in the subantarctic Iles Crozet of the Indian Ocean. Austrodecus is a predominantly 
Subantarctic and Antarctic genus, but with a proposed “‘corridor” of species 
carrying north from New Zealand and the Kermadec Islands to Japan. The closest 
morphological relation to A. palauense, new species, is A. elegans Stock, a species 
known only from the subantarctic Prince Edward Islands. It is less closely related 
to its New Zealand, Kermadec, and Japanese congeners. There is therefore no 
particular pattern to the distribution of the species in this report except perhaps 
a reinforcement to the congeneric corridor theory for Austrodecus. 

The specimens in this report originated from the George Vanderbilt Foundation 
(Stanford University, California) Expedition to the Palau Islands, sponsored by 
the U.S. Office of Naval Research (G.V.F.), a short survey trip to Guam and Palau 
by J. L. Barnard and the author, and from the Bernice P. Bishop Museum (BPBM), 
Honolulu, Hawai. All specimens except the Bishop specimen are deposited in 
the U.S. National Museum of Natural History under the catalog numbers of the 
old U.S. National Museum (USNM). 


Family Ammotheidae 
Achelia nana (Loman) 


Ammothea nana Loman, 1908:60-61, pl. 1, figs. 1-13. 
Achelia nana. —Stock, 1953a:300—301, fig. 14; 1954:97; 1965:14-15, figs. 1-3; 
1968:16; 1974:13-14.—Utinomi, 1971:329-330. 


Material examined. —Saipan, from test block immersed in sea 90 days, 24 Feb 
1950, 1 6 (BPBM). 

Remarks. — There is little difference between this specimen and published figures 
of males. There is a hint of a segmentation line between the first and second trunk 
segments, but none between the second and third. The sixth oviger segment has 
the strong recurved spine mentioned by Stock (1965:15). 

This specimen extends the distribution slightly to the east from the known Indo- 
West-Pacific habitats of Achelia nana. 


Achelia, species indeterminate 


Material examined.— Palau, SE Koror Island, Iwayama Bay, E side of mouth 
of Raki-Swido (Oyster Pass), from sponge in 2 to 15 ft. (0.6—4.6 m), 7°18'57’N, 
134°30'09”E, 22 Oct 1955, G. V. F. sta 220A, 1 juvenile. Palau, Koror Island, 
sea reef inside cove next to cave at SE end of Island, wash of 4 kinds of sponges 
in 0.1 m, 2 Jul 1974, J. L. Barnard, 1 2. 

Remarks.—The juvenile specimen has some characters of Achelia assimilis 
(Haswell), but this variable species demands an adult with a full set of characters 
for assignment. The female from the cave cove is possibly a new species. The 
trunk is semi-triangular in dorsal aspect and has no tubercles except at the an- 
terolateral corners of the cephalic segment. The first coxae have 2 small laterodistal 


700 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 1. Ammothella elegantula, male: a, Trunk, dorsal view; b, Trunk, lateral view, with distal 
chelifore enlargement; c, Palp; d, Third leg, with distal segments enlargement; e, Oviger; f, Strigilis, 
with terminal denticulate spine enlarged. 


tubercles, but the trunk and appendages are otherwise without tubercles. The 
distal 4 palp segments are almost circular in outline and the second segment is 
only twice as long as its diameter. Without a male, I refrain from naming this 
specimen rather than add another name to this confusing group of extremely 
variable species. 


VOLUME 96, NUMBER 4 701 


Ammothella elegantula Stock 
Fig. 1 


Ammothella elegantula Stock, 1968:11-—12, fig. 2a—c. 


Material examined.— Palau, Koror Island, sea reef inside cove next to cave at 
SE end of island, wash of 4 kinds of sponges in 0.1 meter, 2 Jul 1974, J. L. Barnard, 
1 6, 2 larvae. 

Remarks.—This very distinctive species is reported here only for the second 
time. It was described from a juvenile specimen taken in the Straits of Malacca 
in 77 meters. The present material fortunately includes a male from which the 
adult characters may be described. 

The chelae, as in all adults of Ammothella, are reduced to rounded knobs having 
2 short stubs as fingers. The ocular tubercle does not have the 2 posterior tubercles 
of the type-specimen, but has many tiny low papillae over most of its surface. 
The abdomen does not have any of the long setae or tubular spines common to 
species of this genus, but has a few short distal setae. The setae of the lateral 
processes and first coxae are distinctive. They are longer than the segment diameter 
and occur distally on the posterior of the first lateral processes and on the anterior 
and posterior of the remaining 6 lateral processes. Those of the first coxae are 
longer, fully twice the segment diameter or slightly longer, and are 4 in number, 
placed dorsodistally. 

In the adult, palp segments 2 and 4 are subequal and are the longest segments. 
Segments 5 and 6 are twice as long as wide and subequal in length, and the 
terminal 3 segments are small, only slightly longer than wide. 

The male oviger is typical of Ammothella; second segment longest, fourth seg- 
ment slightly shorter, fifth about two-thirds the length of the fourth, the seventh 
with a slight distal apophysis bearing 2 long setae ectally and | denticulate spine 
endally, eighth and ninth with 1 denticulate spine each, and tenth a tiny segment 
bearing 2 larger denticulate spines having 5 to 8 marginal serrations. 

The leg is long and very slender, as described, and has a few very long setae 
per segment, some 4 or 5 times longer than the segment diameter. The femur and 
first tibia are swollen distally, or club-shaped, while the second tibia is cylindrical. 
The cement gland appears as a dorsodistal tube over twice as long as the femur 
diameter. The terminal leg segments agree very much with those of the juvenile 
with 2 heel spines and a similar tarsus spine. The second tibia is slightly shorter 
than the first in the adult and the femur is slightly shorter than either tibia. Most 
other measurements differ only slightly from the juvenile holotype. 

The capture of this species in Palau extends its distribution well out into the 
Western Pacific islands from the Straits of Malacca, and its depth range is extended 
from 77 meters to the littoral. 


Ammothella tippula, new species 
Fig. 2 


Material examined.—Guam, Anae Island, just E in deep channel, wash of 
coralline and other algae from 11 m, 13°21'25”N, 144°38'20”E, C. A. Child, 27 
Jun 1974, 1 éholotype (USNM 195374), 1 2, 1 juvenile paratypes (USNM 195375). 


702 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 2. Ammothella tippula, holotype male: a, Trunk, dorsal view; b, Trunk, lateral view; c, Palp, 
distal 7 segments; d, Third leg; e, Terminal segments of third leg enlarged; f, Oviger; g, Strigilis. Female: 
h, Strigilis. 


VOLUME 96, NUMBER 4 703 


Description. — Very small, holotype leg span slightly more than 5 mm. Trunk 
broad, robust, fully segmented, without median tubercles or spines, armed with 
2 small slender anterolateral tubercles on ocular segment. Lateral processes 1.5 
times longer than wide, separated by half their diameters or less, armed with tiny 
low dorsodistal tubercles bearing papillae and short distal setae on posterior and 
anterior of all lateral processes except first where anterior setae are missing. Ocular 
tubercle tall, over 3 times taller than maximum diameter, swollen at apex with 
large darkly pigmented eyes, capped with tiny papilla. Abdomen a long posterior- 
curing cylinder with median and distal swellings armed with 2 groups of very 
long tubular spines and 2 very long distal setae, 2 short median setae and 2 short 
laterodistal setae. 

Proboscis ovoid, massive, with broad flat lips having ventral cleft just distal to 
marked constriction. 

Chelifores 3-segmented, robust, almost as long as proboscis, second segment 
only slightly longer than first. First segment armed with long dorsal tubular spine 
and 3 lateral and distal setae. Second segment armed with long dorsal tubular 
spine, 1 long seta of equal length just proximal to spine, and 5 long dorsodistal 
and laterodistal setae, each longer than segment diameter. Chela small, ovoid, 
vestigial, with crease for movable finger. 

Palp 9-segmented, slender, segments 3 to 9 increasingly setose distally. Segment 
2 little longer than segment 4, both armed with several lateral and ventrodistal 
setae longer than segment diameter; segments 5 and 6 subequal, over 3 times 
longer than their diameter; segment 7 shortest, less than twice longer than its 
diameter; segment 8 twice its diameter in length; terminal segment slightly less 
than 3 times longer. Most setae longer than segment diameter. 

Oviger 10-segmented, segments 2 and 4 subequal, segment 5 slightly longer. 
Seventh segment swollen distally with apophysis bearing 3 long setae. Seventh, 
eighth and ninth segments armed with single denticulate spine each, bearing many 
marginal denticulations. Terminal segment tiny, wider than long, armed with 2 
denticulate spines larger than preceding ones. 

Third leg: first coxa armed with 2 long tubular spines on first pair of legs, 1 
similar spine on posterior 6 legs, 2 dorsodistal long setae and | lateral short seta 
on anterior and posterior of all first coxae. Coxa 2 with dorsal bulge bearing 1 
long tubular spine and | long seta, several setae distally. Coxa 3 with several 
ventral and ventrodistal setae. Femur shorter than tibiae, armed with 4 lateral 
long tubular spines and 2 long lateral setae, ventrally with 2 short setae, and 
distally with 4 or 5 long setae. Cement gland tube dorsodistal, longer than segment 
diameter, inserted on low tubercle. Tibia 2 slightly longer than tibia 1, both armed 
with 2 dorsal long tubular spines, 2 long setae proximally and | long seta distally 
measuring more than 3 times segment diameter, several short lateral setae and 2 
or 3 tubular spines, and several short ventral setae. Tarsus tiny, armed with 1 
dorsal, 2 lateral, and 1 ventral setae and 1 ventrodistal spine. Propodus moderately 
slender, well curved, without heel but with 3 large heel spines and 5 or 6 small 
sole spines, dorsally armed with several setae longer than segment diameter and 
few short distal setae. Claw moderately long, slightly curved with well curved 
auxiliaries only slightly shorter than main claw. 

Female (paratype) at least 25 percent larger in all measurements except oviger 


704 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


which is smaller than that of male. Oviger segment lengths similar to male but 
strigilis without seventh segment apophysis and setae. Denticulate spines arranged 
2:1:1:2 from seventh to tenth segments. Fewer tubular spines on most appendages. 

Measurements of holotype (n mm).—Trunk length (chelifore insertion of tip 
4th lateral processes), 0.51; trunk width (across 2nd lateral processes), 0.46; pro- 
boscis (laterally), 0.39; abdomen (laterally), 0.38; 3rd leg: coxa 1, 0.13; coxa 2, 
0.23; coxa 3, 0.16; femur, 0.44; tibia 1, 0.48; tibia 2, 0.51; tarsus, 0.07; propodus, 
0.27; claw, 0.11. 

Distribution.—Known only from the type-locality, Anae Island, Guam, in 
JUL saat. 

Etymology—The species name is Latin and refers to a water spider. It is some- 
times spelled tipula. 

Remarks. —There are a number of Ammothella species which share many char- 
acters with A. tippula, all of which appear to form a discrete species group within 
the genus: the appendiculata group. The Pacific members of this group are A. 
appendiculata (Dohrn), A. indica Stock, A. pacifica Hilton, A. schmitti Child, A. 
dawsoni Child and Hedgpeth, 4. spinifera Cole, A. symbius Child, A. setacea 
(Helfer), and A. elegantula Stock. These species are all characterized by having a 
more or less slender appearance, long abdomen, slender legs bearing long setae 
and sometimes long spines, long chelifores with long setae or spines, fairly long 
ocular tubercle, and typical terminal leg segments with long claws. Most of these 
species bear some form of tubular, clubbed, or spatulate spines on the appendages, 
and it is often the presence or absence of these “‘special”’ spines that distinguishes 
a particular species. In relating these species to A. tippula, A. elegantula (reported 
on elsewhere in this report) is probably the most distantly related due to its lack 
of tubular spines, its very long slender proboscis, chelifores and ocular tubercle, 
and other dissimilar arrangements of setae and spines. The very long ocular 
tubercle and slender proboscis also disqualify A. setacea as a close relation. In 
addition, A. setacea has very long tubular spines on the cephalic segment and 
lateral processes, unlike A. tippula. 

The remaining species all have a long curved abdomen bearing long setae, and 
most also have the long abdominal tubular spines as in A. tippula, but A. appen- 
diculata, A. indica, and A. symbius have first scape segments much shorter than 
the second, unlike A. tippula, and have other characters such as spine arrangement 
which disagree with those of the new species. Ammothella spinifera has middorsal 
tubular spines on the trunk (in most but not all specimens) and tall slender lateral 
process tubercles unlike A. tippula. Ammothella dawsoni has many more spines 
on the chelifores and has many bristled or plumose spines on the appendages 
which are not found on A. tippula, while A. schmitti has similar plumose spines 
in a different arrangement, a short ocular tubercle and more robust propodus with 
shorter claws. 

The closest relation to A. tippula is A. pacifica, another Pacific island species. 
The new species differs from A. pacifica in having small tubercles at the antero- 
lateral corners of the ocular tubercle, tiny low lateral process tubercles, three heel 
spines instead of four, an ocular tubercle placed well posterior to the rim of the 
ocular segment instead of adjacent to the rim, and a generally more robust ap- 
pearance. The size of these two species offers further contrast. The leg span of an 
adult male of A. pacifica is 8.24 mm, while that of A. tippula is only 5.26 mm. 


VOLUME 96, NUMBER 4 705 


Ammothella pacifica is known from several localities in the Indo-West Pacific, 
including the Caroline Islands at Ifaluk Atoll, and is very close to the new species 
in spination and the segment lengths of its appendages. 


Tanystylum rehderi Child 
Tanystylum rehderi Child, 1970:302-—306, fig. 5. 


Material examined.— Palau, 8 mi NW of Koror, on inner margin of barrier 
reef, 7°24'30’N, 134°21'18”E, from octocoral Clavularia sp., with sand and coral 
in 1-2 m, 19 Jul 1955, G.V.F. sta 25-865, 2 6 with eggs, 1 2, 1 juvenile. 

Remarks.— These specimens differ little from the type-specimens, 2 males from 
Bora Bora and Moorea, Society Islands. The ocular tip tubercle is a larger cone 
in the Palau specimens. Most of the other tubercles and segment lengths agree 
with the types. 

The female was unknown until this Palau record. The lateral process and the 
first coxa tubercles are reduced in the female and those of the first coxa anterior 
are not bifurcate as in the male. The female oviger is typical of the genus and has 
spines which are only distally denticulate on the terminal 3 segments in the ratio 
eee 

This species is related to Janystylum acuminatum, as discussed by Child (1970: 
305), but the shape of the proboscis is quite different in 7. rehderi, being a low- 
shouldered, long-necked bottle shape, while in 7. acuminatum, the proboscis 1s 
like a broad oval flask having a narrow and shorter neck. 

The Palau specimens extend the distribution of 7. rehderi from the Society 
Islands westward to the western Caroline Islands and increase its depth range to 
2 meters from the intertidal of the type-specimens. 


Family Phoxichilidiidae 


Anoplodactylus chamorrus, new species 
Fig. 3 


Material examined.—Guam, Anae Island, just E in deep channel, wash of 
coralline and other algae from 11 m, 13°21'25"N, 144°38'20’E, C. A. Child, 27 
Jun 1974, 1 6 holotype (USNM 195376). 

Description.— Very small, leg span slightly less than 4.5 mm. Trunk unseg- 
mented, robust, broadly oval in dorsal aspect. Lateral processes touching proxi- 
mally, slightly separated distally, little longer than their diameters, each armed 
with single low, broad, dorsodistal tubercle, glabrous. Ocular tubercle short, cy- 
lindrical proximally, a rounded cone distally, with large darkly-pigmented eyes 
at tubercle midlength. Neck short, narrow, glabrous. Abdomen erect, cylindrical, 
tapering distally to rounded point, armed with 2 posterolateral short setae. 

Proboscis ovoid with slight proximal constriction, tapering distally to flat lips. 

Palps represented by tiny buds on anterior of first lateral processes. 

Chelifores robust, overhanging proboscis, scape armed with several short lateral 
and distal setae. Chela palm longer than fingers, oval, armed with 3 or 4 endal 
and ectal setae. Fingers short, well curved, without teeth, armed with 2 or 3 short 
setae ectally on movable finger. 

Oviger moderately long, first segment broad, distal half with anterior expansion, 


706 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 3. Anoplodactylus chamorrus, holotype male: a, Trunk, dorsal view; b, Trunk, lateral view, 
with oviger attached; c, Third leg, with cement gland enlarged; d, Chelifore. 


third segment longest, about 1.2 times as long as second segment, both second 
and third armed with several short setae. Strigilis with few setae; fourth with 2 
or 3 ectal setae, fifth with 6 or 7 recurved short setae, terminal segment a curved 
cone with 3 or 4 short setae. 

Leg moderately short, robust, armed with 1 dorsodistal seta almost twice as 
long as segment diameter of each major segment, few short setae on all segments. 
Femur longest segment, single cement gland a small oval terminating in slender 


VOLUME 96, NUMBER 4 707 


distally-pointing tube about 0.3 as long as segment diameter. Second coxa with 
small ventral sexual spur not extending beyond distal end of segment, terminating 
in tiny sex pore. Tarsus semi-rectangular, without long ventrodistal extension, 
armed with 2 ventral setae. Propodus robust, with marked heel bearing | heel 
spine and 2 smaller spines, sole straight, armed with 5 curved spines, several tiny 
setae and small propodal lamina only 0.25 as long as sole. Claw massive, mod- 
erately curved, auxiliaries absent. 

Measurements of holotype (in mm).—Trunk length (chelifore insertion to tip 
4th lateral processes) 0.6; trunk width (across | st lateral processes), 0.45; proboscis 
(lateral), 0.38; abdomen (lateral), 0.15; third leg: coxa 1, 0.12; coxa 2, 0.28: coxa 
3, 0.13; femur, 0.35; tibia 1, 0.3; tibia 2, 0.22; tarsus, 0.08; propodus, 0.29; claw, 
0.19. 

Distribution.—Known only from the type-locality, Anae Island, Guam, in 11 
meters. 

Etymology. Named for the ancient settlers of Guam and the Mariana Islands, 
the Chamorros, whose friendliness is immediately evident to modern travellers 
in these islands. 

Remarks.—This tiny species resembles several other Anoplodactylus species 
having crowded lateral processes, short neck, and short robust legs. These are; A. 
anarthrus Loman, A. aragaoi Sawaya, A. arescus d-B. R. Marcus, A. compactus 
(Hilton), A. derjugini Losina-Losinsky, A. haswelli (Flynn), A. marcusi Mello 
Leitao, A. minusculus Clark, A. tarsalis Stock, and A. viridintestinalis (Cole), 
several of which were described in the now junior synonym Halosoma. 

The new species is perhaps least related to A. minusculus, which has extremely 
short oviger segments, a long propodal lamina, and large chela fingers with teeth. 
The presence or absence of chela teeth is not always a good diagnostic character. 
Sometimes with a good series of one species, some specimens may not have chela 
teeth while others will have them. Anoplodactylus haswelli has three heel spines, 
no propodal lamina, a triangular-shaped trunk, auxiliary claws and chela finger 
teeth, while A. aragaoi has slightly separated lateral processes, longer appendage 
segments, a longer neck, and a longer cement gland tube. This new species is 
unlike A. tarsalis and A. arescus, both of which have a projecting tarsus of a long 
triangular-shape and differing leg segment lengths. 

Anoplodactylus chamorrus is more closely related to A. anarthrus, except that 
the proboscis of the latter is smaller and of a different shape, the ocular tubercle 
is a more rectangular shape, and the cement gland is placed more proximally on 
the femur. The propodus of A. viridintestinalis has a lamina over the full sole and 
has longer appendage segments than 4. chamorrus. The figures of A. marcusi 
(Mello Leitao, 1949:167-173, figs. 1-4) are inconclusive in detail, but they show 
a smaller proboscis, shorter leg segments, and different oviger segment lengths 
than those of A. chamorrus. 

The new species appears to be closest to A. derjugini and A. compactus. The 
differences are that these two species have a non-tubular cement gland, full pro- 
podal lamina, auxiliary claws, and in A. compactus, larger lateral process tubercles, 
and with A. derjugini, a differently shaped chela with longer fingers, different 
ocular tubercle length, two heel spines, and many more setae on the terminal 
Oviger segment. The combination of characters in 4. chamorrus are thus unlike 
those of any known species. 


708 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Anoplodactylus pycnosoma (Helfer) 


Peritrachia pycnosoma Helfer, 1938:176-177, fig. 7. 
Halosoma pycnosoma.—Marcus 1940:45—46. 
Anoplodactylus pycnosoma.—Stock 1953:41, fig. 5; 1954:75-77, fig. 33; 1974:16; 

1975:132.—Utinomi 1971:326.—Child 1975:20. 

Material examined.— Palau, Ngaremediu District, Urukthapel Island, fringing 
coral reef around first rock cape W of sand beach, 7°15'18”N, 134°26'48”E, on 
red algae in 0-2 m, 31 Jul 1955, G.V.F. sta 53-980, 1 2. Palau, Urukthapel Island, 
small bay at E end of N shore, around remains of stone pier, 7°15'57’N, 
134°26'55”E, in 0.6-1.3 m, 2 Nov 1955, G.V.F. sta 258, 1 °. 

Remarks.— These two females differ only slightly from the published figures for 
this species. The lateral processes of the Palau specimens are further apart and 
the propodus is slightly longer, but the specimens agree in other characters in- 
cluding the brown lines at the trunk segmentation and lateral process-first coxa 
segmentation. 

The Palau Islands are added to the Indo-West Pacific distribution for this 
species. It is apparently confined to littoral depths. 


Anoplodactylus, species indeterminate 


Material examined.— Palau, SE Koror Island, Iwayama Bay, E side of mouth 
of Raki-Swido (Oyster Pass), on sponge in 2-15 ft (0.6-—4.6 m), 22 Oct 1955, 
G.V.F. sta 220A, 7°18'57’N, 134°30'09’E, 1 larva. Palau, SE Koror Island, sea 
reef inside cove next to cave, 7°18'34”N, 134°30'35”E, wash of 4 kinds of sponges 
in 0.1 m, J. L. Barnard, 2 Jul 1974, 3 larvae. 

Remarks.— There are at least two species represented in the three larvae col- 
lected by Barnard, but none of these specimens is of sufficient age to be determined. 


Family Callipallenidae 
Callipallene novaezealandiae (Thomson) 


Pallene novae-zealandiae Thomson, 1884:246—247, pl. 14, figs. 1-4. 
Callipallene sp. cf. C. novaezealandiae.—Child 1982:277 [literature]. 


Material examined. — Palau, unnamed islet between Amoi and Eil Malk islands, 
7°11'35”N, 134°23'00’E, from hydroid-ascidian-sponge complex in 2 m, C. A. 
Child, 30 Jun 1974, 2 6 with eggs, 1 2, 6 juveniles. Palau, SE Koror Island, sea 
reef inside cove next to cave, 7°18'34”N, 134°30'35”E, wash of 4 kinds of sponges 
in 0.1 m, 2 Jul 1974, J. L. Barnard, 1 °. 

Remarks.—These specimens agree in all respects with those reported (Child 
1982:277) from Enewetak Atoll. One of the males has more strigilis denticulate 
spines than the Enewetak specimens, which decreases the gap of differences be- 
tween these specimens and Thomson’s original description of the species. I there- 
fore propose to assign both the Enewetak and the Palau specimens to Thomson’s 
species. 

The Enewetak and Palau records extend the previously known distribution of 
this species from east Africa, Australia and New Zealand, to the northern hemi- 
sphere in Micronesia. 


VOLUME 96, NUMBER 4 709 


Seguapallene Pushkin, 1975 


This genus was designated (Pushkin 1975:1404—1405, fig. 2) to contain a species 
of pycnogonid from the subantarctic Iles Crozet: Seguapallene insignatus Pushkin. 
This species is without palps, has well developed 2-segmented chelifores with 
regular teeth on the fingers, auxiliary claws, and an oviger with strigilis claw. The 
principal difference between this species and the new one described herein is teeth 
on the strigilis claw in the latter, lacking in Pushkin’s species. 


Seguapallene micronesica, new species 
Fig. 4 


Material examined. — Palau, SE Koror Island, sea reef inside cove next to cave, 
7°18'34’N, 134°30'35”E, wash of 4 kinds of sponges in 0.1 m, 2 Jul 1974, J. L. 
Barnard, | 6 holotype (USNM 195379), 3 2, 2 juveniles, paratypes (USNM 195380). 

Description.— Moderately small, male leg span less than 9 mm. Trunk fully 
segmented, stout, without median tubercles, glabrous. Lateral processes short, not 
longer than wide, separated by half their diameter or less, glabrous. Neck short, 
only as long as width of oviger bases. Ocular tubercle implanted over oviger bases, 
short, rounded, only as tall as basal diameter, with tiny posteromedian bump and 
small lateral “horn” tubercles slightly longer than their basal widths. Eyes large, 
darkly pigmented. Abdomen moderately short, not extending beyond first coxae 
tips of fourth pair of legs, cylindrical anteriorly, tapering posteriorly, armed with 
pair of posterolateral setae. 

Proboscis short, ovoid, tapering to small flat lips, without fringe of oral setae. 

Chelifore 2-segmented, scape as long as proboscis, armed with 2 short ectal 
setae, 2 midlateral setae longer than segment diameter and 2 or 3 dorsodistal setae 
as long as segment diameter. Chela palm shorter than fingers, armed with several 
long dorsal and lateral setae. Fingers straight, curved only at tips, armed with 
short broad teeth, 11 on immovable finger and 10 on movable finger. 

Oviger fourth and fifth segments long, fourth 7 times longer than its diameter, 
fifth over 8 times its diameter, armed with several ectal setae slightly longer than 
segment diameter. Fifth segment with distal apophysis shorter than segment di- 
ameter, armed with distal seta. Four strigilis segments cylindrical, each segment 
shorter than preceding, armed with denticulate spines having 4 lobes on each 
margin, arranged in the formula 5:5:4:5, well curved terminal claw bearing 5 large 
teeth. 

Third leg; first coxa armed with anterodistal seta as long as coxa diameter, coxa 
2 with 4 lateral setae longer than segment diameter, coxa 3 with several shorter 
ventral and ventrodistal setae. Femur and tibiae with several long lateral setae 
and from | to 4 longer dorsal setae, longer than twice segment diameter. Cement 
glands a series of tiny ventral pores along most of femur length. Tibia 2 longest 
segment. Tarsus quadrilateral, short, armed with 2 ventral setae longer than tarsus 
diameter. Propodus slender, moderately curved, without heel or heel spines, armed 
with 5 or 6 sole setae as long as segment diameter and several dorsal and distal 
setae, dorsodistal 2 over twice propodus diameter. Main claw short, not as long 
as segment diameter, well curved. Auxiliary claws more than twice main claw 
length, strongly curved. 

Female (paratype) slightly larger in all measurements. Oviger segments 4 and 


710 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 4. Seguapallene micronesica, holotype male: a, Trunk, dorsal view; b, Trunk, lateral view; c, 
Proboscis, ventral view; d, Chela; e, Third leg; f, Terminal segments of third leg enlarged; g, Oviger; 
h, Strigilis terminal segment enlarged. 


5 shorter, length only 3 times diameter, apophysis absent, strigilis with fewer 
denticulate spines and only 3 teeth on terminal claw. Main claw of propodus 
slightly shorter than in male. 

Measurements of holotype (in mm).— Trunk length (chelifore insertion to tip 
4th lateral processes), 0.95; trunk width, 0.41; proboscis (lateral), 0.38; abdomen 
(lateral), 0.23; third leg, coxa 1, 0.23; coxa 2, 0.46; coxa 3, 0.2; femur, 0.83; tibia 
1, 0.96; tibia 2, 1.02; tarsus, 0.09; propodus, 0.36; main claw, 0.07; auxiliary 
claws, 0.15. 


VOLUME 96, NUMBER 4 711 


Distribution.— Known only from the type-locality, Koror Island, Palau Islands, 
in the intertidal. 

Etymology.—Named for Micronesia, of which Palau and the Caroline islands 
form a major part. 

Remarks.—The only other known species in this genus, Seguapallene insignatus 
Pushkin (1975:1404—1405, fig. 2), 1s quite different from this new species. Push- 
kin’s species has pointed tubercles on the lateral processes, a much longer ab- 
domen, a major ventral spine on the tarsus and three major heel spines, a large 
normal-size claw with two smaller auxiliaries, and marked differences in the 
oviger. Pushkin (1975:fig. 2f, g) illustrates what appears to be a female oviger 
while labeling it that of a male. The fourth and fifth segments are much shorter 
than those of S. micronesica and lack the fifth segment apophysis. His figure ‘“‘g”’ 
is of the oviger terminal segments and shows a smooth claw without teeth. The 
denticulate spines differ greatly from the new species in having two long proximal 
serrations and many short distal serrations per side, similar to those common to 
the genus Callipallene. The denticulate spines of S. micronesica are more like 
those of many of the ammotheids. 

The discovery of S. micronesica gives the genus an extraordinary distribution 
pattern. Seguapallene insignatus was described from specimens found in the cold 
Iles Crozet of the Subantarctic in 3 to 30 meters. The new species, found in 
tropical Palau, shows that the genus is not confined to particular thermal habitats, 
but that it is perhaps only confined to littoral and sublittoral habitats. There is a 
somewhat similar distribution pattern in the genus Austrodecus (see following 
species). This genus has a predominantly Antarctic and Subantarctic distribution, 
but several species are known to occur from New Zealand and the tropical west 
Pacific to Japan. 


Family Austrodecidae 


Austrodecus palauense, new species 
Fig. 5 


Material examined.— Palau, SE Koror Island, sea reef inside cove next to cave, 
7°18'34’N, 134°30'35”E, wash of 4 kinds of sponges in 0.1 m, 2 Jul 1974, J. L. 
Barnard, 1 2 juvenile holotype (USNM 195377), 1 juvenile paratype (USNM 
195378). 

Description. — Holotype very tiny, leg span only 2.05 mm. Trunk slender, elon- 
gate, completely segmented, with 4 slender median tubercles shorter than trunk 
diameter, anterior tubercle longest, those posterior increasingly shorter. Lateral 
processes separated by their diameters or slightly more, armed with tiny latero- 
distal tubercles on posterior of first pair, anterior and posterior of second and 
third pairs, and absent from last pair. Ocular tubercle an obliquely anterior- 
pointing tapered cone, placed at extreme anterior of neck, bulging distally with 
well pigmented small eyes, over 3 times longer than its maximum diameter, apex 
armed with 2 tiny lateral papillae. Proboscis typical of the genus, long, tubular 
with distal annulations. Abdomen a long cylinder tapering distally, extending to 
distal tip of second coxae of fourth pair of legs, armed with 2 small lateral subapical 
setae. 

Palp 5-segmented, second segment longest, armed with few short setae, 1 longer 


712 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 5. Austrodecus palauense, holotype juvenile female; a, Trunk, dorsal view; b, Trunk, lateral 
view; c, Palp; d, Palp terminal segments enlarged; e, Third leg; f, Mid-dorsal tubercle of ocular segment 
enlarged. 


ventrodistal seta and | slender dorsodistal tubercle not as long as segment di- 
ameter. Fourth segment curved, 0.6 as long as second, armed with several short 
setae, 3 long dorsal and ventral setae, and 3 stout curved spines dorsodistally. 
Fifth segment subrectangular with ventral tubercle giving appearance of slight 
bifurcation of segment, armed distally with dense setae longer than segment di- 
ameter. 

Ovigers represented only by rudimentary buds ventrally on cephalic segment. 

First coxae with dorsolateral tubercles slightly shorter than segment diameter, 
some with distal setae, arranged with | posteriorly on coxae of first pair of legs, 
2 on second and third pair, and | anteriorly on fourth pair of legs. Third coxae 
with small dorsal tubercle not as long as segment diameter. Major leg segments 
armed with few short setae and single large dorsodistal setae longer than 3 times 
segment diameter. Propodus moderately curved, as long as femur, armed with 2 
or 3 short sole setae, 2 dorsodistal setae and well curved claw slightly more than 
0.3 propodus length. Sexual characters undeveloped. 


VOLUME 96, NUMBER 4 713 


Measurements of holotype (4n mm).—Trunk length (palp insertion to tip 4th 
lateral processes), 0.59; trunk width (across 2nd lateral processes), 0.25; proboscis, 
0.63; abdomen, 0.16; ocular tubercle, 0.22; 3rd leg, coxa 1, 0.05; coxa 2, 0.07; 
coxa 3, 0.06; femur, 0.16; tibia 1, 0.15; tibia 2, 0.17; tarsus, 0.03; propodus, 0.15; 
claw, 0.06. 

Distribution.— Known only from Koror Island, Palau Islands, in 0.1 meter. 

Etymology.—Named for the type-locality, the Palau Islands. 

Remarks.—The discovery of this new species in a genus with predominantly 
Antarctic distribution tends to reinforce the theory that there is a western Pacific 
“corridor” of species from New Zealand to Japan, to account for the known 
temperate and tropical species development in this area. There is no known 
counterpart for this corridor along the eastern coast of South America or the east 
African coast. Several (at least five) species are known from New Zealand, two 
from the Kermadec Islands, the new species reported on herein from the Palau 
Islands, and the single known species from Japan all point toward this northern 
corridor. It may be possible, in future collecting, to discover other species in this 
corridor in the New Hebrides, Solomons, New Guinea, Marianas, and the Bonin 
Islands, as many species of Austrodecus appear to be endemic or at least are 
confined to restricted areas or corridors of distribution (see Stock 1957:figs. 7, 8, 
12). 

Austrodecus palauense appears closely related to A. elegans, a species from near 
the Subantarctic Prince Edward Islands. Both have well separated lateral processes, 
mid-dorsal trunk tubercles, slender laterodistal coxa | tubercles, and a very similar 
abdomen. The ocular tubercle of A. elegans is much longer and anterior-pointing, 
the leg segments are much longer, and it has auxiliary claws. How the new species 
will appear as an adult is conjecture, but it would already have developed auxilary 
claws at this stage of growth if it were to have them. It 1s not likely that the lateral 
processes would be spaced closer together in the adult than in the juvenile (or 
sub-adult). In all other known species of Austrodecus, the lateral processes are 
more closely spaced than in 4. palauense. 


Acknowledgments 


I wish to thank the late Dr. Dennis M. Devaney for loaning the Bishop Museum 
material, and my colleagues, J. L. Barnard, for his infectuous collecting enthu- 
siasm, and T. E. Bowman, both of the Department of Invertebrate Zoology (Crus- 
tacea), National Museum of Natural History, for reviewing the manuscript. 


Literature Cited 


Child, C. A. 1970. Pycnogonida of the Smithsonian-Bredin Pacific Expedition, 1957.— Proceedings 
of the Biological Society of Washington 83(27):287-308. 
1975. Pycnogonida of Western Australia.—Smithsonian Contributions to Zoology 190:1- 

28. 

. 1982. Pycnogonida of the Western Pacific Islands I. The Marshall Islands. — Proceedings of 

the Biological Society of Washington 95(2):270-281. 

Helfer, H. 1938. Einige neue Pantopoden aus der Sammlung des Zoologischen Museums in Berlin. — 
Sitzungsberichte Gesellschaft Naturforschungs Freunde (Berling) 1937:162-185. 

Loman, J.C.C. 1908. Die Pantopoden der Siboga-Expedition.-Siboga Expedition, Monographie 40: 
1-88. 

Marcus, E. 1940. Os Pantopoda brasileiros e os demais Sul-Americanos.—Boletim Faculdade de 
Filosofia, Ciéncias e Letras da Universidade de Sao Paulo, series 19 (Zoologia) 4:3-179. 


714 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Mello Leitao, A. de. 1949. Nova Espécie de Halosoma Cole, 1904 (Pantopoda, Phoxichilidiidae 
Sars, 1891).—Anais da Academia Brasileira de Ciencias 21(2):167-173. 

Pushkin, A. F. 1975. [New Species of Callipallenidae (Pantopoda) from the Subantarctic.]—Zool- 
ogicheskii Zhurnal 54(9):1402-1406, 2 figures. [In Russian plus English Summary]. 

Stock, J. H. 1953. Re-description of some of Helfer’s pycnogonid type-specimens. — Beaufortia 4(35): 

33-45. 

. 1953a. Contribution to the knowledge of the pycnogonid fauna of the East Indian Archipelago. 

Biological Results of the Snellius Expedition 17.—Temminckia 9:276-313. 

——. 1954. Pycnogonida from Indo-West-Pacific, Australia, and New Zealand waters. — Viden- 
skabelige Meddelser fra Dansk Naturhistorisk Forening (Copenhagen) 116:1—-168. 

—. 1957. The Pycnogonid family Austrodecidae.—Beaufortia 6(68):1-81. 

——. 1965. Pycnogonida from the southwestern Indian Ocean.—Beaufortia 13(151):13-33. 

. 1968. Pycnogonida collected by the “Galathea”’ and ““Anton Bruun”’ in the Indian and Pacific 

Oceans. — Videnskabelige Meddeleser fra Dansk Naturhistorisk Forening (Copenhagen) 131:7— 

65. 

1974. Medio- and infralittoral Pycnogonida collected during the I. I. O. E. near Landbase 
on Nossi-Be, Madagascar. — Bulletin of the Zoological Museum, University of Amsterdam 4(3): 
11-22. 

1975. Infralittoral Pycnogonida from Tanzania.—Travais du Muséum d’Histoire Naturelle 
“Gr. Antipa’’, Bucharest 16:127-134. 

Thomson, G. M. 1884. On the New Zealand Pycnogonida with descriptions of new species. — 
Transactions and Proceedings of the New Zealand Institute 16(1883):242—248. 

Utinomi, H. 1971. Records of Pycnogonida from shallow waters of Japan.— Publications of the Seto 
Marine Biological Laboratory 18(5)317-347. 


Department of Invertebrate Zoology, Smithsonian Institution, Washington, D.C., 
20560. 


PROC. BIOL. SOC. WASH. 
96(4), 1983, pp. 715-718 


RECOGNITION OF TWO SPECIES OF 
DOUBLE-LINED MACKERELS 
(GRAMMATORCYNUS: SCOMBRIDAE) 


Bruce B. Collette 


Abstract.— Grammatorcynus has been considered to be monotypic by recent 
authors. Electrophoretic work by A. D. Lewis indicates the presence of two species 
in Australia. These are identified herein as: (1) G. bilineatus (Riippell, 1836); type- 
locality: Red Sea; “‘scad’’; a wide-spread species (Red Sea to Tonga); with many 
gill rakers (19-24), large eye (7-9% of fork length), and smaller maximum size 
(60 cm FL, 3 kg); and (2) G. bicarinatus (Quoy and Gaimard, 1844); type-locality: 
Shark Bay, Western Australia; ““shark mackerel’’; limited to the northern coasts 
of Australia and the Gulf of Papua; few gill rakers (12-15), small eye (3-5% of 
FL), and larger maximum size (110 cm FL, 13.5 kg). 


Most recent authors (e.g., Silas 1963; Collette 1979) have considered Gram- 
matorcynus to be monotypic. In correspondence with me and in his doctoral 
dissertation, Lewis (1981) presented electrophoretic evidence that there are two 
species in Australia: a small species, the “scad,” that is unspotted, and a larger 
one, the “‘shark mackerel,” that frequently has dark spots on its lower sides. This 
brief paper presents morphological evidence for recognition of two species: the 
widely distributed scad, G. bilineatus (Riippell) and the shark mackerel G. bicar- 
inatus (Quoy and Gaimard), restricted to Australia and southern New Guinea. 
This evidence supports inclusion of both species in the FAO World Catalogue of 
scombrids (Collette and Nauen, in press). A revision of Grammatorcynus with 
discussion of its relationships to other scombrids is in progress. 


Grammatorcynus bilineatus (Riippell, 1836) Scad 


Fig. 1A 
Thynnus bilineatus Riippell, 1836:39—40, pl. 12, fig. 2 (original description, Red 
Sea). 
Grammatorcynus bilineatus Gill, 1862:125 (T. bilineatus type-species of new ge- 
nus). 


Nesogrammus piersoni Evermann & Seale, 1907: 61-62, pl. 1, fig. 3 (original 
description; Bulan, Sorsogon Province, Luzon, Philippine Is.). 


Diagnosis.—A species of Grammatorcynus with many gill rakers, (3-5) + 1 + 
(13-18) = 19-24 on the first arch (Table 1); a large eye, 7-9% of FL (see Fig. 2); 
matures at a small size, about 430 mm FL; maximum size about 600 mm FL, 3 
kg. Seldom with dark spots on the lower sides of the body. 

Range.— Widespread in the Indo-West Pacific. Based on the literature and 
material examined, known from the Red Sea, Andaman Sea, East Indies, Phil- 
ippines, Ryukyu Islands, New Guinea (New Britain, New Ireland, New Hanover, 
and the Louisiade Archipelago), Australia (Scott Reef off northern Western Aus- 


716 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


B 


Fig. 1. Two species of Grammatorcynus. A, G. bilineatus (from Evermann and Seale 1907, fig. 3, 
holotype of Nesogrammus piersoni, 372 mm FL, Philippine Islands); B, G. bicarinatus (from Mc- 
Culloch 1915, pl. 1, fig. 1, 925 mm FL, New South Wales, Australia). 


tralia, eastern Queensland), the Solomon Islands, New Caledonia, the Caroline 
Islands, Marshall Islands, Fiji, and Tonga. 

Material examined.—52 specimens (23.5-575 mm FL) from 36 lots: Red Sea 
(13 specimens including Senckenburg Museum 2755, the stuffed holotype of Thyn- 
nus bilineatus), Andaman Sea (4), Celebes (2), Philippines (5 including USNM 
55899, the holotype of Nesogrammus piersoni), New Guinea (7), Australia (7), 
Solomons (1), Carolines (3), Marshalls (8), and Fiji (2). 


Grammatorcynus bicarinatus (Quoy and Gaimard, 1824) Shark Mackerel 
Fig. 1B 


Thynnus bicarinatus Quoy and Gaimard, 1824:357, pl. 61, fig. 1 (original de- 
scription; Baie des Chiens-Marins = Shark Bay, W. Australia). 

Grammatorcynus bicarinatus McCulloch, 1915:266—269, pl. 1, fig. 1 (description; 
off Cook Is., near Tweed River Heads, New South Wales; 925 mm FL, 18.75 
Ibs.). 


Diagnosis.—A species of Grammatorcynus with few gill rakers, (1-2) + 1 + 
(10-12) = 12-15 on first arch (Table 1); a small eye, 3—4% of FL (see Fig. 2); 
reaches large size, probably 1100 mm FL, 13.5 kg. Frequently has dark spots on 
the lower sides of the body (Fig. 1B). 


VOLUME 96, NUMBER 4 717 


28 > 
26 
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50 60 70 80 90 100 110 120 130 140 150 160 


HEAD LENGTH (mm) 


Fig. 2. Regression of orbit length on head length in two species of Grammatorcynus. Regression 
equation for G. bilineatus (n = 44), Y = 2.923 + 0.189X; for G. bicarinatus (n = 9), Y = 3.593 + 
0.144X. Probability that regressions are the same (by ANCOVA), P < 0.0000001; that the slopes are 
equal, P = 0.0032; and that the intercepts are equal, P < 0.0000001. 


Table 1.—Total number of gill rakers in the species of Grammatorcynus. 
Species and area 12 13 14 15 16 7 18 19 20 Dl 22: 23 24 N xX 


G. bilineatus 


Red Sea 1 © 2 2) l WA Qie7 
Andaman Sea 3 1 4 22.3 
Celebes Islands 2 2 20.0 
Philippine 

Islands 1 — Dior al 4 20.8 
New Guinea 1 3 i 1 6 20.5 
Australia 1 2 ltl 5 20.4 
Solomon Islands ] it Dil. 
Caroline Islands 1 1 1 3 B20 
Marshall Islands 1 1 6 8 20.6 
Fiji Islands 1 1 2 240.5 

Total 4 10 19 8 5 i ay Dilsh 


G. bicarinatus 1 = § 2 9 14.0 


718 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Range.—Positively known only from the northern coasts of Australia with 
occasional stragglers south to 30°S on both east (Cook Is., N.S.W.) and west (Shark 
Bay, W.A.) coasts and in the Gulf of Papua off southern New Guinea (A. D. 
Lewis, pers. comm.). 

Remarks.—The original description of bicarinatus is not detailed enough to tell 
which species is involved, the figure is poor, and there is no type-material. I use 
the name because the shark mackerel extends further south on the coasts of 
Australia than does the scad, to Cook Island, N.S.W. on the east (McCulloch 
1915) and to Exmouth Gulf, W.A. (USNM uncat.) and, presumably Shark Bay 
on the west. 

Material examined.—9 specimens (300-825 mm FL) from 8 lots: Western 
Australia (4); Queensland (5). 


Acknowledgments 


I thank Dr. A. D. Lewis (Department of Agriculture and Fisheries, Fiji) for 
informing me of this problem and for providing frozen specimens of both species 
for me to dissect. Sally Rothwell spent the month of February 1983 working in 
my laboratory on Grammatorcynus on a Careers in Biology Program from Colgate 
University. Her efforts have made early completion of this preliminary paper 
possible. I thank the curators of the collections housing material of Grammator- 
cynus for permission to study their material and I will acknowledge their help 
individually in the revision of the genus. Ruth Gibbons ran the analysis of co- 
variance and prepared the figures. Drafts of the manuscripts were read by Robert 
H. Gibbs, Jr., A. D. Lewis, Victor G. Springer, and Austin B. Williams. 


Literature Cited 


Collette, B. B. 1979. Adaptations and systematics of the mackerels and tunas. — Jn The physiological 
ecology of tunas, Gary Sharp and Andrew Dizon, eds.; Academic Press, N.Y., pp. 739. 
Collette, B. B., and C. Nauen. (In press). Scombrids of the world. FAO Species Catalogue, vol. 3, 
Food and Agriculture Organization of the United Nations Fisheries Synopsis No. 125. 
Evermann, B. W.,and A. Seale. 1907. Fishes of the Philippine Islands. — Bulletin of the [U.S.] Bureau 
of Fisheries 26:49-110. 

Gill, T. N. 1862. On the limits and arrangement of the family of scombroids.— Proceedings of the 
Academy of Natural Sciences of Philadelphia 14:124—127. 

Lewis, A. D. 1981. Population genetics, ecology and systematics of Indo-Australian scombrid fishes, 
with particular reference to skipjack tuna (Katsuwonus pelamis). Australian National University, 
Ph.D. thesis. 

McCulloch, A.R. 1915. Notes on, and descriptions of Australian fishes. — Proceedings of the Linnean 
Society of New South Wales 40:259-277. 

Quoy, J. R. C., and J. P. Gaimard. 1824. Voyage autour du Monde, ... Exécuté sur les corvettes 
de S.M. l’Uranie et la Physicienne, pendant les années 1817, 1818, 1819 et 1820. 3, Zoologie, 
712 pp. 

Riippell, E. 1836. Neue Wirbelthiere zu der Fauna von Abyssinien gehorig. Fische des rothen Meeres, 
6:29-52. Frankfurt am Main. 

Silas, E.G. 1963. Synopsis of biological data on double-lined mackerel Grammatorcynus bicarinatus 
(Quoy and Gaimard) (Indo-Pacific). — Food and Agriculture Organization of the United Nations 
Fishery Report No. 6, 2:811-833. 


National Marine Fisheries Service Systematics Laboratory, National Museum 
of Natural History, Washington, D.C. 20560. 


PROC. BIOL. SOC. WASH. 
96(4), 1983, pp. 719-724 


TWO NEW SPECIES OF CORAL TOADFISHES, 
FAMILY BATRACHOIDIDAE, GENUS SANOPUS, 
FROM YUCATAN, MEXICO, AND BELIZE 


Bruce B. Collette 


Abstract. — Sanopus reticulatus, new species, is described from three specimens 
collected at Progreso, Yucatan, Mexico. It differs from the other species of Sanopus 
in its reticulate body pattern and shorter distance from snout to second dorsal fin 
origin (403-419 versus 420-487 thousandths of standard length). It is most closely 
related to S. barbatus and S. johnsoni, species with mottled bellies, branched chin 
barbels, and high numbers of fin rays and vertebrae. Sanopus greenfieldorum, new 
species, is described from three specimens collected at Carrie Bow Cay, Belize. It 
is most closely related to S. astrifer but has light lines on its head instead of light 
spots. 


While examining fishes in the Zoological Museum of the University of Hamburg 
(ZMH) in September 1982, I was surprised to discover a fine specimen of toadfish 
collected in Yucatan in 1893 that was identified as Opsanus tau. It was clearly a 
specimen of Sanopus with a distinctive pattern and was from the Gulf of Mexico, 
beyond the known range of the genus (Cozumel Is., Quintana Roo, south to 
Panama); I suspected it represented an undescribed species. Subsequently, two 
additional specimens of this species were located at the Museum of Comparative 
Zoology (MCZ) at Harvard that had been misidentified as Opsanus pardus by 
Barbour and Cole (1906). A second new species was discovered first by David 
W. and Teresa A. Greenfield on the barrier reef near Carrie Bow Cay, Belize, in 
1974 about the time my (1974) revision of Sanopus was published. Another 
specimen of this species was collected at Carrie Bow Cay in 1980, and a juvenile 
in 1983. 

The purpose of this paper is to describe these two new species and to compare 
them with the four known species in the genus: S. barbatus (Meek and Hildebrand), 
S. astrifer (Robins and Starck), S. splendidus Collette, Starck, and Phillips, and 
S. johnsoni Collette and Starck (see Collette 1974). 

Counts and measurements follow those given in my review of Sanopus. Tables 
of meristic data presented there are not repeated here. Material of the four pre- 
viously treated species housed at the National Museum of Natural History (USNM) 
was compared with the new species. Specimens of S. astrifer and S. barbatus from 
the Field Museum of Natural History in Chicago (FMNH) collected subsequent 
to 1974 were examined to increase the sample size of comparative material. 


Sanopus reticulatus, new species 
Figs. 1A—B 
Opsanus pardus non Goode and Bean, Barbour and Cole, 1906:159 (two speci- 


mens from Progreso). 


Holotype.—ZMH 20930, male, 236 mm SL, Mexico, Yucatan, Progreso; Weiss, 
25 Jan 1893. 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


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VOLUME 96, NUMBER 4 721 


Fig. 2. Diagrammatic anterodorsal views of heads of two species of Sanopus. A, S. greenfieldorum 
(holotype, USNM 213555, 280 mm SL, Carrie Bow Cay, Belize); B, S. astrifer (USNM 209720, 245 
mm SL, Glovers Reef, Belize). 


Paratypes.—MCZ 32889 (233 mm SL) and USNM 258111 (225 mm), Mexico, 
Yucatan, Progreso, Leon J. Cole, early 1904. 

Diagnosis. — Differs from the other five species of Sanopus in having a reticulate 
pattern on the body and head (Fig. 1A). Similar to S. barbatus and S. johnsoni 
and different from S. astrifer, S. greenfieldorum, and S. splendidus in having 
branched chin barbels. Maxillary barbel branched as in the first two species but 
more deeply divided than in either of them (Fig. 1B—D). Distance from tip of 
snout to origin of second dorsal fin (Table 1) shorter (403-419 thousandths of 
SL, x 413.7) than in the other species (420-487, means 432.5—451.0). No cirri 
present in interorbital region as are present in S. johnsoni (Fig. 1D). 

Description. — Dorsal fin rays III, 31-32; anal fin rays 25-26; pectoral fin rays 
19-20; vertebrae 11 + (27-29) = 38-40; upper lateral line papillae 34-36; lower 
lateral line papillae 30-32; dentary teeth 18-19; palatine teeth 9-14; total vo- 
merine teeth 8—10; and premaxillary teeth 34—41 in one or two rows on each jaw. 
Body proportions given in Table 1. 

Etymology.—Named in reference to its diagnostic color pattern. 

Biology.—The holotype is a male with well-developed gonads. Both paratypes 
are immature. The holotype had a specimen of majid crab in its stomach, Mac- 
rocoeloma trispinosum (Latreille). There was a snail in the intestine of one of the 
paratypes. 


Sanopus greenfieldorum, new species 
Fig. 2A 


Holotype. —USNM 213555, male, about 280 mm SL, Belize, 2 mi. S of Carrie 
Bow Cay; D. W. and T. Greenfield, G74-21; 24 Jul 1974. 

Paratypes.—FMNH 94575, ripe female, about 270 mm SL, Belize, between 
Carrie Bow Cay and South Water Cay; E. Reynaud; 10 Mar 1980. USNM 261601, 
29.3 mm, Carrie Bow Cay; G. Hendler; 2 Apr 1983. 

Diagnosis.—Similar to S. astrifer in having a dark body with prominent light 
markings. Differs from S. astrifer in having many light lines on head (Fig. 2A) 


22 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Table 1.—Morphometric comparison (in thousandths of standard length) between adults of the six 
species of Sanopus. 


S. reticulatus S. barbatus 
(n = 3) (n = 12) S. johnsoni 
Character Range x Range x 
SL (mm) DQM A. -— BBN BNF 370 8§=©309..5 253 
Head length 326 363 340.00 341 307 BUSS) 352 
Head width 245 303 265.3 295 322 310.9 295 
Orbital 40 45 43.3 34 52 43.2 42 
Interorbital 61 77 71.0 85 108 95.3 89 
Snout-2 dorsal fin 
origin 403 419 413.7 427 471 447.5 439 
Snout—anal fin origin 587 592 589.3 572 647 # 604.9 581 
Pectoral fin length 162 183 172.3 157 199 172.3 175 
Pelvic fin length 132 157 140.7 131 159 146.6 153 
S. greenfieldorum S. astrifer S. splendidus 
(n = 2) (n = 4) (n= 
Range x “Range Range x 
SL (mm) QQ 230 2ISO 233 2637 9 24453 9) 219, ee SeO 
Head length 361 415 388.0 354 368 360.5 338 356 ©=—. 345..8 
Head width 318 356 8 §©=—6337.0 323 333 327.3 290 314 299°5 
Orbital 41 41 41.0 41 50 46.0 40 52 46.2 
Interorbital 92 113 102.5 84 97 90.3 69 V7 73.5 
Snout-—2 dorsal fin 
origin 439 575 507.0 420 487 444.8 423 439 432.5 
Snout—anal fin origin 467 659 563.0 570 597 #£2585.5 583 604 594.0 
Pectoral fin length 190 202 196.0 193 219 206.8 158 203 178.6 


Pelvic fin length 157 166 161.5 144 180 162.0 154 203 170.0 


instead of small light spots (Fig. 2B). About 4 continuous light lines between eyes, 
several more on top of head posterior to these, and 6—8 more lines radiating out 
from lower part of orbit. Eye slightly smaller than in S. astrifer (41 thousandths 
of SL, 99-115 thousandths of head length compared to 41-50 and 116-139). 

Description. — Dorsal fin rays II, 30-32; anal fin rays 24—25; pectoral fin rays 
22; vertebrae (11-12) + (26-27) = 37-39; upper lateral line papillae 36—40; lower 
lateral line papillae 30-33; dentary teeth 22-28; palatine teeth 14—16; total vo- 
merine teeth 10-16; and premaxillary teeth 31-35 in one or two rows on each 
jaw. Body proportions given in Table 1; both adult type specimens bent, therefore, 
measurements of SL, snout to second dorsal, and snout to anal could not be made 
with accuracy. 

Underwater photographs of a specimen larger than the types taken by James 
Bohnsack show a pattern similar to that in Fig. 2A but with a few more and 
slightly wider lines on the head. The lines are white on a gray-black background. 

Etymology.—Named for David W. and Teresa Arambula Greenfield who col- 
lected the holotype, suspected it was undescribed, and sent it to me for exami- 
nation. The Greenfields have been working actively on the fish fauna of Belize 
for more than a decade and described a new toadfish, Triathalassothia gloverensis, 
from Glovers Reef in 1973 (but it also occurs at Carrie Bow Cay). 


VOLUME 96, NUMBER 4 723 


Biology.— The holotype was taken by spear from a depression in the sand under 
a large coral head (Vontastrea) in one meter of water on the reef flat behind the 
barrier reef. The fish was facing out so only the front of the head was seen. This 
is the same type of habitat in which S. astrifer has been taken at Glovers Reef 
(D. W. Greenfield, pers. comm.). The adult paratype was also taken by spear from 
under a dead coral clump in slightly deeper water on the reef flat behind the barrier 
reef between Carrie Bow Cay and South Water Cay. James Bohnsack (pers. comm.) 
photographed an S. greenfieldorum in the spur and groove zone of the fore reef 
at Carrie Bow Cay at a depth of about 6 m at about 9:30 PM during the last week 
of April 1982. It was under a coral head facing out with a little more of the fish 
showing than in Fig. 2A. The juvenile paratype was taken with rotenone at 24.4 
m over an area of Montastrea on the fore reef crest. 

The holotype is a male with slightly developed testes. The gut contents consist 
of fragments of two specimens of the portunid crab, Portunus vocans (A. Milne 
Edwards), one specimen of a majid crab, Mithrax, probably M. pleuracanthus 
Stimpson, and a few fish bones. The adult paratype has two large ovaries that 
occupy much of the body cavity. There are 191 eggs in the left ovary, 178 in the 
right. The eggs are mostly 5 or 6 mm in diameter. The teeth of the adult paratype 
are pink and the stomach and intestine were filled with spines and broken pieces 
of the test of Diadema antillarum Philippi. Two crabs were among the gut con- 
tents, a female Portunus vocans and a chela and carpus from a xanthid crab. There 
were also fragments of a skull and a few vertebrae of a small fish. The juvenile 
paratype contained the carapace of an alpheid shrimp, an undescribed species 
of the isopod Stenetrium, and a small snail, Tricolia cf. affinis (C. B. Adams) of 
the family Phasianellidae. 


Discussion 


In my 1974 diagnosis of Sanopus, I noted that it lacked the discrete glands on 
the posterior surface of the pectoral fin between the bases of the upper fin rays 
that are present in Opsanus. I did not point out that there is a well-developed 
glandular area inside the gill cavity anterior to the pectoral girdle. This shared 
specialization is characteristic of adults and subadults of all six species of Sanopus. 

Description of two more species of Sanopus raises the toadfish fauna of the 
western Atlantic to 30 (including three freshwater species in rivers draining into 
the western Atlantic) in 7 genera (Collette and Russo 1981: table 13). The New 
World batrachoid fauna now comprises 43 out of a total of about 66 species in 
the family, confirming the New World as the center of diversity for the family. 

Crabs are an important component of the diet of the two new species of Sanopus 
as they are in other species of toadfishes (Collette and Russo 1981). Diadema was 
present in the gut of the adult paratype of S. greenfieldorum, in the recently 
collected specimen of S. astrifer from Glovers Reef (FMNH 91034) and in the 
holotype of S. johnsoni (Collette 1974). Randall et al. (1964) reported 15 species 
from 7 other families as Diadema predators. 

Egg number and size in S. greenfieldorum are comparable to other toadfishes, 
i.e., relatively few large eggs. Females of 9 species of Batrachoides ranging from 
106-352 mm SL had 88-588 eggs, 3—6 mm in diameter (Collette and Russo 1981: 
table 10) compared to S. greenfieldorum, 270 mm SL with 369 eggs, 5-6 mm in 
diameter. 


724 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Comparative Material Examined 


See Collette (1974) for data on Sanopus examined up to that time. Additional 
material, all from FMNH, is as follows. S. astrifer: 91034 (1, 263), Belize, Glovers 
Reef; 20 June 1978. S. barbatus: 91030(1, 330, Honduras, Brus Lagoon; 7 May 
1975. 91031(2, 295-330), Honduras, Big Hog Is.; 21 May 1975. 91032(1, 328), 
Honduras, Big Hog Is.; 20 May 1975. 91033(1, 300), Honduras, Little Hog Is.; 18 
May 1975. 


Acknowledgments 


I thank Dr. H. Wilkens (ZMH) and Mr. Karsten Hartel (MCZ) for loaning me 
the specimens of S. reticulatus. Dr. Karel Liem (MCZ) and Mr. Hartel kindly 
permitted me to retain one MCZ specimen for the USNM collections. Dr. David 
W. Greenfield collected the holotype of S. greenfieldorum, called my attention to 
it, and permitted me to retain it for the USNM collections. Dr. Donald Stewart 
(FMNH) loaned material of S. greenfieldorum, S. astrifer, and S. barbatus. The 
drawings are by Keiko Hiratsuka Moore. Dr. James Bohnsack provided me with 
color underwater photographs of a specimen from a night dive at Carrie Bow 
Cay. Radiographs of the types were made by Ruth Gibbons. Dr. Austin B. Wil- 
liams identified crustaceans and Dr. Richard S. Houbrick identified the snail in 
the stomachs of the types. Dr. David W. Greenfield and Dr. Williams read drafts 
of the manuscript. 


Literature Cited 


Barbour, Thomas, and Leon J. Cole. 1906. Reptilia, Amphibia, and Pisces. 7m Vertebrata from 
Yucatan.— Bulletin of the Museum of Comparative Zoology 50:146-159. 

Collette, Bruce B. 1974. A review of the coral toadfishes of the genus Sanopus with descriptions of 

two new species from Cozumel Island, Mexico. — Proceedings of the Biological Society of Wash- 

ington 87:185-—204. 

, and Joseph L. Russo. 1981. A revision of the scaly toadfishes, genus Batrachoides, with 

descriptions of two new species from the Eastern Pacific.— Bulletin of Marine Science 31(2): 

197-233. 

Greenfield, David W., and Terry Greenfield. 1973. Triathalassothia gloverensis, a new species of 
toadfish from Belize (=British Honduras) with remarks on the genus.—Copeia 1973(3):560- 
565. 

Randall, John E., Robert E. Schroeder, and Walter A. Starck. 1964. Notes on the biology of the 
echinoid Diadema antillarum.—Caribbean Journal of Science 4(2 + 3):421-433. 


National Marine Fisheries Service Systematics Laboratory, National Museum 
of Natural History, Washington, D.C., 20560. 


PROC. BIOL. SOC. WASH. 
96(4), 1983, pp. 725-757 


A REVISION OF THE SEGUENZIACEA VERRILL, 
1884 (GASTROPODA: PROSOBRANCHIA). I. 
SUMMARY AND EVALUATION OF THE SUPERFAMILY 


James F. Quinn, Jr. 


Abstract.—A summary of the superfamily Seguenziacea is compiled from the 
literature and unpublished observations, and a complete bibliography is presented. 
A taxonomic resumé of seguenziacean genera is given. A preliminary classification 
of the superfamily includes 74 nominal species and subspecies in seven genera 
and four species-groups within Seguenzia. The known characters of the shells and 
anatomy are reviewed. The superfamily Seguenziacea is shown to be distinct from 
any other known archaeogastropod or mesogastropod superfamily. This super- 
family is characterized by: nacreous shells of archaeogastropod ultrastructure, 
often complexly sculptured with 0-3 (usually 2 or 3) labral sinuses; modified 
rhipidoglossate radula (formula 12-4.1.1.1.4-12); paucispiral corneous operculum; 
epipodial tentacles; monopectinate ctenidium; long intestine with an anterior loop; 
specialized structures in the reproductive tract (e.g., a well developed penis); and 
modification of the mantle edge to form distinct incurrent and excurrent siphons. 
Contents of the intestine of Seguenzia sp. cf. S. eritima Verrill indicate that 
Seguenzia is a detritivore. 

Ancistrobasis is known from the Eocene, Pliocene, and Recent; Seguenzia occurs 
from the Miocene to the Recent; all other genera are unknown as fossils. Although 
probably derived from the Trochacea, no direct link with any known fossil or 
living prosobranch group has yet been established. The superfamily Seguenziacea 
is here considered to be an isolated offshoot of the Trochacea, independently 
acquiring advanced anatomical features of a mesogastropod nature as a conse- 
quence of extremely small body size and in response to a deep-water habitat. 


The enigmatic superfamily Seguenziacea comprises a group of very small (usu- 
ally 5 mm or less in height), trochoid-like prosobranchs of world-wide distribution. 
Although a few species have been recorded from outer continental shelf or abyssal 
plain depths, by far the majority of known species have been described from the 
continental slopes. Because of their deep-water habitats, information concerning 
seguenziacean species has been largely confined to original species descriptions 
and records published in reports of major national exploratory expeditions. Very 
few species are represented in collections by large series of specimens, and even 
fewer have been collected alive. Asa result, the taxonomy and systematic position 
of the Seguenziacea have remained in states of confusion and debate. As a first 
step towards resolving the status and relationships of this group, a review of all 
information in the literature is presented, augmented by original observations. 

The observations summarized here suggest that seguenziaceans are archaeo- 
gastropods which have acquired anatomical specializations more typical of a 
mesogastropod organization. Archaeogastropod affinity is indicated by the na- 
creous shell, protoconch, epipodial tentacles, anterior loop of the intestine, and 


726 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


modified rhipidoglossate radula. Features in common principally with mesogas- 
tropods include a monopectinate ctenidium, specialized reproductive system in- 
cluding development of a penis, and functional modification of the mantle edge 
to form distinct incurrent and excurrent siphons. A similar combination of char- 
acteristics is not found in any other known gastropod group, and the argument 
for a separate superfamily, the Seguenziacea, as suggested by Keen (1971), Golikov 
and Starobogatov (1975), Goryachev (1979), Quinn (1981, 1983a, b), McLean 
(1981), and Marshall (in press), is supported. Although a probable origin within 
the Trochacea is postulated, phylogenetic relationships of the Seguenziacea are 
still unclear. 

Seventy-four nominal species are here assigned to seven genera plus four species- 
groups within Seguenzia within the Seguenziacea and a preliminary classification 
is presented (Table 1). No attempt to determine species synonymies was made 
since that task is properly the province of a full monographic review. Such a 
monograph will be published as time and material permit. However, consideration 
of shell morphology and radular characters indicates that several rather discrete 
species-groups exist within Seguenzia sensu lato. These species-groups, not for- 
mally defined here, are referred to in the text and Table 1 as Seguenzia Groups 
I-IV, designations used here for convenience of discussion and as an indication 
of areas requiring close attention in future studies. Species names are used in the 
text without citation of date of description as this information may be found in 
Table 1. A synopsis of the history of the supraspecific taxa is presented and a 
complete bibliography is included in the Literature Cited. 


Taxonomic Resumé 


Seguenzia was proposed almost simultaneously in 1876 by the British mala- 
cologist J. G. Jeffreys and the Italian paleontologist G. Seguenza. It appears that 
Jeffreys’ paper (15 June 1876) was published prior to Seguenza’s (May—June 1876), 
but there is still some doubt as I have been unable to determine an exact date of 
publication for Seguenza’s paper; therefore, pending acquisition of further infor- 
mation, Jeffreys is here considered the author of Seguenzia. Although Jeffreys 
and Seguenza agreed on the name for the genus, they disagreed strongly on its 
supposed systematic position. Seguenza (1876) assigned the group to the Trochi- 
dae. Jeffreys (1876), on the other hand, thought that Seguenzia was most closely 
related to the Solariidae (=Architectonicidae) and later emphatically restated that 
opinion (Jeffreys 1879) after Watson (1879a) erected the second seguenziacean 
genus, Basilissa, and allocated it, along with Seguenzia, to the Trochidae on the 
basis of the nacreous shells. 

Verrill (1884) was the first to examine the radula of Seguenzia, which he de- 
scribed as taenioglossate. On that evidence he erected the family Seguenziidae, 
included both Seguenzia and Basilissa, and placed the family near Aporrhais 
(Strombidae). Based on specimens from the collections of the Blake Expedition, 
Dall (1889b) defined Ancistrobasis as a subgenus of Basilissa, and placed both in 
the Trochidae. Dall assigned Fluxina discula to the Solariidae (=Architectonici- 
dae) and placed the Seguenziidae near the Trichotropidae, approximating Tryon’s 
(1887) allocation of Seguenzia. Dautzenberg and Fischer (1897a) defined Basi- 


VOLUME 96, NUMBER 4 VAT 


lissopsis for a small, distinctively shaped seguenziacean which they included in 
the Trochidae. 

Schepman (1908, 1909) contributed perhaps the most information of any author 
to date to the understanding of the Seguenziacea. In his 1908 report, Schepman 
described and illustrated radulae of Basilissa and his new genus, Guttula, and 
although he retained these two genera in the Trochidae, he commented on the 
uniqueness of their radular formulae. In the second part of his report (1909), he 
illustrated the radula of Seguenzia melvilli, the first accurate description and clear 
illustration of a radula of Seguenzia. He speculated that Seguenzia would even- 
tually be recognized as belonging to a rhipidoglossate group also encompassing 
Basilissa, although he followed Tryon (1887), Dall (1889b), and explicitly Pel- 
seneer (1906), in placing the Seguenziidae near the Trichotropidae. Dall (1925) 
included his new subgenus, Orectospira, in Basilissa, but Habe (1955a, b) has 
shown that Orectospira is turritellid rather than seguenziid. 

Reflecting the hiatus in research on the deep sea, it was not until 1971 that 
another generic taxon was established. Thelyssa Bayer, 1971, was erected for a 
species described from University of Miami collections of deep-water Caribbean 
molluscs. Thelyssa is very similar to, and may eventually be considered a subgenus 
of, Basilissa. The most recently defined genus, Mioseguenzia Nordsieck, 1973, 
was introduced for Janthina cimbrica Sorgenfrei, a Miocene fossil from Denmark, 
and two new taxa, M. cimbrica recens and M. conica. Dr. Philippe Bouchet (in 
litt.) has informed me that his examination of Nordsieck’s specimens revealed 
them to be larval shells of the Cypraeacea. From Sorgenfrei’s illustration, it appears 
that M. cimbrica is also a larval form, thus excluding Mioseguenzia from the 
Seguenziacea. 

The genus F/uxina Dall, 1881, has been used for a number of seguenziaceans 
(Dall 1889b; Schepman 1909; Bayer 1971) and recently included in a list of 
seguenziacean genera (Boss 1982); however, Merrill (1970a) has shown that the 
type-species of Fluxina, F. brunnea Dall, is a Calliostoma (Trochidae). Fluxiella 
(Okutani, 1968, 1974) is a nomen nudum, and F. vitrea Okutani, is here assigned 
to Seguenzia Group IV. Quinn (1983b) recently erected Carenzia for the Se- 
guenzia carinata species-group, and Marshall (in press) is treating the species- 
groups referred to herein as Seguenzia Groups II and IV, as well as three other 
new genera. 


Shell Morphology 


Species of the Seguenziacea, although not uncommon in collections of deep- 
water molluscs, are rarely represented by large suites of shells, and even fewer by 
live-collected specimens. Resulting identifications and classifications have been 
based almost solely on shell characters. The taxonomic value of shell characters 
has not been critically evaluated, and authors have labored under considerable 
uncertainty as to which variations are merely intraspecific, and which actually 
represent specific differences. This inadequacy is perhaps best illustrated by the 
following comments of W. H. Dall (1889b:269): “In examining the specimens of 
Seguenzia ...I1 find myself in a dilemma. Either each separate individual is to 
be regarded as a species or the variability of the shells is very great. Persistent 


728 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Table 1.—Preliminary classification of the Seguenziacea. 


Seguenziacea Vermill, 1884 


Seguenziidae Verrill, 1884 


Seguenzia Jeffreys, 1876 


Type-species.— Seguenzia formosa Jeffreys, 1876; by monotypy. 


Group I (SeguenZia s.s.) 


S. 
S. 
S. 
TS. 


+S. monocingulata var. elegans Seguenza, 1876 


+S. 


—- 
ANANRNDRNNNDNDHDHHDHDDHDDNNDYDDY 


~~ 


nn 


TS. 


formosa Jeffreys, 1876 

formosa var. lineata Watson, 1879 
formosa var. nitida Verrill, 1884 
monocingulata Seguenza, 1876 


monocingulata var. elata Seguenza, 1876 
eritima Verrill, 1884 

elegans Jeffreys, 1885 

elegans vat. bicarinata Locard, 1898 
occidentalis Dall, 1908 

stephanica Dall, 1908 

costulifera Schepman, 1909 
dautzenbergi Schepman, 1909 
melvillii Schepman, 1909 

certoma Dall, 1919 

giovia Dall, 1919 

cervola Dall, 1919 


. caliana Dall, 1919 
. antarctica Thiele, 1925 


sumatrensis Thiele, 1925 


. orientalis Thiele, 1925 

. floridana Dall, 1927 

. hapala Woodring, 1928 
. hosyu Habe, 1953 

. louiseae Clarke, 1961 

. fatigans Barnard, 1963 
. soyoae (Okutani, 1964) 
. mirabilis Okutani, 1964 


nipponica Okutani, 1964 


. megaloconcha Rokop, 1972 


donaldi Ladd, 1982 


Group II 


S. 
S! 
S. 
iS: 
S. 


S. 


lonica Watson, 1879 
lampra (Watson, 1879) 
polita Verco, 1906 

sykesi Schepman, 1909 
cazioti Dautzenberg, 1925 
rushi Dall, 1927 


Group III 


S. 
S. 


siberutensis Thiele, 1925 
simplex Barnard, 1963 


Group IV 


S. 
2S. 


S. 


discula (Dall, 1889) 

dalliana (Melvill and Standen, 
1903) 

marginata (Schepman, 1909) 


_ NE Atlantic Ocean 


W Atlantic Ocean 

NW Atlantic Ocean 

Pliocene, Italy 

Pliocene, Italy 

Pliocene, Italy 

NW Atlantic Ocean 

NE Atlantic Ocean 

NE Atlantic Ocean 

NE Pacific Ocean 

NE Pacific Ocean 

Celebes, Indonesia 

Celebes, Indonesia 

Celebes, Indonesia 

NE Pacific Ocean 

NE Pacific Ocean 

NE Pacific Ocean 

NE Pacific Ocean 

S of Kerguelen, Southern Ocean 
SW of Sumatra, Indian Ocean 
E of Kenya, Indian Ocean 

W Atlantic Ocean 
Miocene-Recent, W Atlantic Ocean 
Tosa Bay, Shikoku, Japan 

SE Atlantic Ocean 

S of Madagascar, SW Indian Ocean 
Off Torishima Is., Japan 

Off Aoga-shima Is., Japan 

Sea of Enshu-Nada, Japan 

NE Pacific Ocean 

Pleistocene, New Hebrides 


NW Atlantic Ocean 

E of Japan, NW Pacific Ocean 
S of South Australia 

Banda Sea, Indonesia 

NE Atlantic Ocean 

NW Atlantic Ocean 


SW of Sumatra, Indian Ocean 
Off Cape Point, South Africa 


NW Atlantic Ocean 
N Indian Ocean 


Banda Sea, Indonesia 


VOLUME 96, NUMBER 4 


Table 1.— Continued. 


S. trochiformis (Schepman, 1909) 
S. stenomphala (Melvill, 1910) 

S. gelida (Barnard, 1963) 

2S. solarium (Barnard, 1963) 

S. vitrea (Okutani, 1968) 


Carenzia Quinn, 1983 


V2) 


Ceram Sea, Indonesia 

N Indian Ocean 

W of Cape Point, South Africa 

S of Madagascar, SW Indian Ocean 
S of Boso Peninsula, Honshu, Japan 


Type-species. — Seguenzia carinata Jeffreys, 1877; by original designation. 


C. carinata (Jeffreys, 1877) 
C. trispinosa (Watson, 1879) 
C. inermis (Quinn, 1983) 


Basilissa Watson, 1879 


W and NE Atlantic Ocean 
W Atlantic Ocean 
NE Pacific Ocean 


Type-species. — Basilissa superba Watson, 1879; by subsequent designation, Cossmann, 1888. 


. alta Watson, 1879 

. alta var. oxytoma Watson, 1879 
. simplex Watson, 1879 

. munda Watson, 1879 

. superba Watson, 1879 

. sibogae Schepman, 1908 

. watsoni Dall, 1927 


Ancistrobasis Dall, 1889 


Bas awsadh 


NW Atlantic Ocean 

NW Atlantic Ocean 

SW Atlantic Ocean 

E Atlantic Ocean 

N of Australia, SW Pacific Ocean 
Celebes, Indonesia 

NW Atlantic 


Type-species.— Basilissa costulata Watson, 1879; by subsequent designation, Dall, 1927. 


+A. reticulata (Philippi, 1844) 


A. costulata (Watson, 1879) 
A. lusitanica (Fischer, 1882) 
A. depressa Dall, 1889 
+A. radialis (Tate, 1890) 
+A. cossmanni (Tate, 1894) 
A. compsa Melvill, 1904 
2+A. bilix (Hedley, 1905) 
A. bombax (Cotton and Godfrey, 1938) 
+A. pacifica Ladd, 1970 


Basilissopsis Dautzenberg and Fisher, 1897 


Pliocene, Italy; 7Recent, NE 
Atlantic 

NW Atlantic Ocean 

NE Atlantic Ocean 

NW Atlantic Ocean 

Eocene of South Australia 

Eocene of South Australia 

N Indian Ocean 

E of Australia, SW Pacific Ocean 

S of Australia 

Eocene of Tonga, SW Pacific Ocean 


Type-species. — Basilissopsis watsoni Dautzenberg and Fischer, 1897; by monotypy. 


B. oxytropis (Watson, 1879) 

B. watsoni Dautzenberg and Fisher, 
1897 

B. rhyssa (Dall, 1927) 


Thelyssa Bayer, 1971 


S Atlantic Ocean 
NE Atlantic Ocean 


NW Atlantic Ocean 


Type-species. — Thelyssa callisto Bayer, 1971; by original designation. 


T. callisto Bayer, 1971 
Guttula Schepman, 1908 


NW Atlantic Ocean 


Type-species.— Guttula sibogae Schepman, 1908; by monotypy. 


G. sibogae Schepman, 1980 
G. blanda Barnard, 1963 
G. galatheae Knudsen, 1964 


+ = species described from fossil material. 
? = provisional placement. 


N of New Guinea, W Pacific Ocean 
Off Cape Point, S. Africa 
Kermadec Trench, SW Pacific Ocean 


730 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


study of the specimens has convinced me that the latter is the true solution, and 
that the most evident characters, such as the umbilicus (in some adult specimens) 
may be present or absent; that the number of spiral threads, their strength and 
sharpness on the basal disk, are entirely inconstant, and, while in the typical 
formosa the ridge next to the suture is waved or granulate, in many it is perfectly 
plain.” 

Indeed, this is the impression obtained when only a few specimens are examined, 
especially within Seguenzia Group I. However, my examination and comparison 
of a large number of specimens and species indicate that shell characters are not 
as variable as thought; indeed, several appear to be of considerable importance 
in separating species. 

Shell shape.—Shell shapes may be trochoid, turbinate, conoid, or lenticular. 
Outlines of some major species-groups are given in Figs. 1-14. All seguenziacean 
groups except Seguenzia Group III, Ancistrobasis, and Guttula are distinctly car- 
inate, at least at the periphery. Carenzia and Basilissopsis often have a mid-whorl 
ridge or angulation marking the abapical edge of the posterior labral sinus (Figs. 
4, 12), which may approach the strength of the corresponding carina of Seguenzia 
Groups I and II (Figs. 1-3, 5, 6). The latter two species-groups, in addition to 
peripheral and medial carinae, have a basal, and often a subsutural, carina. Major 
intraspecific variation in shell shape is conferred by changes in height-width ratio 
with a concomitant increase or decrease in relative spire height. 

Sculpture. — Guttula and Seguenzia Groups II and IV, and Carenzia are almost 
devoid of sculpture, having at most fine spiral striae (Figs. 4, 7, 10, 11, 14). 
Basilissa, Thelyssa, and Basilissopsis usually have collabral growth lines slightly 
thickened at regular intervals, producing weak, sigmoid, axial riblets, often crossed 
by fine spiral threads (Figs. 9, 12, 13). Continuation of axial sculpture onto the 
peripheral carina often produces a crenulated or scalloped periphery. Seguenzia 
Group II species usually have rather strong spiral basal cords in addition to carinae 
and fine spiral striae (Fig. 3). 

Only Seguenzia Group I and Ancistrobasis exhibit strong sculptural patterns 
(Figs. 1, 2, 5, 6, 8). Ancistrobasis has reticulate sculpture of subequal axial and 
spiral cords with nodules produced at the intersections, and the base bears strong, 
obscurely nodulous or undulate spiral cords. The intricate sculpture of Seguenzia 
Group I is by far the most striking of all seguenziaceans. In addition to the three 
or four spiral carinae previously described, the shell bears strong basal cords, fine 
spiral threads between carinae, and collabral riblets. The abrupt changes in di- 


— 


Figs. 1-12. Seguenziacean shells: 1, 2, 5, 6, Seguwenzia Group I; 3, S. Group III; 4, Carenzia; 7, S. 
Group IV; 8, Ancistrobasis; 9, Thelyssa; 10, 11, Guttula; 12, Basilissopsis. 1, Seguenzia n. sp., Phil- 
ippines, USNM, SEM, 20x: 2, S. hapala, off West Florida, FDNR, SEM, 33.5; 3, S. rushi, off 
Puerto Rico, USNM, SEM, 13.4: 4, Carenzia carinata, Straits of Florida, UMML, SEM, 13.4~; 5, 
Seguenzia sp. cf. S. elegans, off Yucatan, UMML, SEM, 13.4; 6, S. lineata, off Yucatan, UMML, 
SEM, 20x; 7, S. siberutensis, Philippines, USNM, SEM, 13.4: 8, Ancistrobasis n. sp., off West 
Florida, FDNR, SEM, 13.4; 9, Thelyssa callisto, W of Great Inagua Is., Bahamas, USNM, height 
5.8 mm; 10, 11, Guttula sibogae (from Schepman 1908, pl. II, fig. 7); 12, Basilissopsis watsoni (from 
Dautzenberg 1927, pl. VI, fig. 36). (USNM = U.S. National Museum of Natural History; FDNR = 
Florida Department of Natural Resources, Marine Research Laboratory; UMML = Rosenstiel School 
of Marine and Atmospheric Science, University of Miami.) 


VOLUME 96, NUMBER 4 


ene a Oe 


pene wae soe "GD, 5 


epanetnintn, 


732 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Figs. 13, 14. Seguenziacean shells: 13, Basilissa alta, Straits of Florida, UMML, SEM, 6.5 x; 14, 
Seguenzia Group IV, Seguenzia discula, Straits of Florida, UMML, SEM, 6.5 x. (UMML = Rosenstiel 
School of Marine and Atmospheric Science, University of Miami). 


rection of the collabral lirae reflect the positions of the three labral sinuses char- 
acteristic of this group. 

Labral sinuses.— Reminiscent of many Pleurotomariacea and Turridae, almost 
all Seguenziacea are characterized by the presence of usually two, often three, 
sinuses in the outer lip. Guttula alone exhibits an entire lip (Fig. 10). All other 
groups have a shallow to deep subsutural (=anal) sinus and a shallow sinus in the 
peripheral part of the basal lip. A third, very narrow sinus occurs in the antero- 
lateral part of the lip of Seguenzia Group I, corresponding to the peripheral carina, 
and has been reported to occur in Seguenzia Group II (Watson 1879a; 1886). In 
groups with a strong, flange-like peripheral carina (Seguenzia Group IV and Car- 
enzia, Basilissa, Basilissopsis, and Thelyssa), the anterolateral sinus is represented 
by a channel corresponding to the carina. The outer lip descends from the suture, 
defining the edge of the subsutural sinus which may be J- (Figs. 15, 17), reversed 
L- (Fig. 16), or V- or U-shaped (Fig. 18), then abruptly swings forward perpen- 
dicular to the axis of coiling (as far as 4 whorl in some species of Seguenzia 
Group I), retreats to the anterolateral sinus, advances again for a short distance, 
retreats again to form the basal sinus, and finally arcs forward slightly to the 
columellar region. In undamaged specimens of species of Seguenzia so far ex- 
amined, the edges of the three sinuses are usually distinctly flared and often 
strongly produced into a spout-like process, especially the anal sinus. The “sinus” 
at the base of the columella may be more an artifact of development of the 
columellar tooth rather than primarily of functional significance, although there 
is a shallow, papillate embayment of the mantle edge in Seguenzia sp. cf. S. 
eritima (see Anatomy section). The basal sinus is analogous to the anterior (in- 
halent) siphonal canal of many higher gastropods, and the subsutural sinus cor- 
responds to the anal, or excurrent, sinus of many prosobranchs, most notably the 
pleurotomariaceans and the Turridae. The subsutural sinus apparently appears 
immediately after termination of the protoconch (Figs. 25—29 herein; see also 
Bandel 1979, pl. 1, Figs. 2, 4). The slightly sinuous or straight subsutural riblets 
descend almost perpendicularly before abruptly swinging forward to become con- 
fluent with the mid-whorl carina. This configuration is very similar to that seen 
in adult shells (Figs. 15—17), and seems to contradict Bandel’s statement that no 
subsutural sinus is present prior to the third teleoconch whorl (Bandel 1979:52). 


VOLUME 96, NUMBER 4 733 


Figs. 15-18. SEM micrographs of surface sculpture of Segwenzia and Carenzia species, 134x (s = 
suture): 15, S. hapala; 16, Seguenzia n. sp.; 17, S. lineata; 18, C. carinata. 


Columellar tooth.—A\though the columellae of several seguenziacean groups 
end in blunt, obscure denticles, only Seguenzia Groups I and II, and some An- 
cistrobasis species, have prominent tooth development. The teeth in these groups 
are basically of three types. The Type I tooth, present in Ancistrobasis and a few 
Seguenzia Group I species (Figs. 19, 20), appears as a low to moderate ridge or 
swelling midway down the columella. This ridge may represent the true termi- 
nation of the columella, with the vertical part of the basal lip thickened to form 
an effective continuation of the columellar structure. The Type II, or patulous, 
tooth (Fig. 21) occurs in Seguenzia Group II and occasionally in Group I. It forms 
a very strong, tongue-like projection extending into the aperture and terminating 
a strongly concave columella. In some species the projection is not so pronounced 


734 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Figs. 19-24. SEM micrographs of apertures of Seguenzia and Ancistrobasis: 19, S. hapala, 50x; 
20, Ancistrobasis n. sp., 25; 21, S. floridana, 25x; 22, Seguenzia sp. 50x; 23, S. lineata, 50x; 24, 
Seguenzia n. sp., 50x. 


(Figs. 22, 23). The basal lip descends from the outer edge of the tooth well back 
from the tip (Fig. 21). The Type III tooth (Fig. 24) is also strongly projecting, but 
forms a fairly acute tooth rather than a broad shelf. In this type, the basal lip 
joins along the inner side of the tip. Although usually easily distinguishable in 
specimens in which they are fully developed, teeth of Types IJ and III are often 
indistinguishable during ontogeny. In addition, the tooth may appear at different 
times within the same species, or even within the same population. This usually 
presents little problem in identification; however, it does pose problems in making 
accurate and consistent measurements of shell height, since the collumellar tooth 
is more often preserved than any part of the fragile basal lip. 

Protoconch.— The seguenziacean protoconch is perhaps the most conservative 
shell feature within the group, varying only in size and relative prominence. It is 
very similar to the trochacean protoconch, consisting of about one whorl, some- 
times smooth, but usually sculptured with microscopic granules that usually co- 
alesce into irregular ridges, and ending in a slightly thickened rim (Bandel 1979; 
Figs. 25-30 herein). Bandel (1979) also showed that the mineralogical ultrastruc- 
ture of the protoconch was typically archaeogastropodan (see discussion of shell 
structure below). Size (between 275 um and 400 um in all specimens so far 
examined) and morphology of the seguenziacean protoconch indicate direct or 
lecithotrophic development (see Bouchet 1976; Bouchet and Warén 1979). Size 


VOLUME 96, NUMBER 4 735 


Figs. 25-30. SEM micrographs of protoconchs of Seguenzia, Carenzia, Ancistrobasis, and Basi- 
lissa, 100 x: 25, S. hapala;, 26, Seguenzia sp. cf. S. elegans; 27, Ancistrobasis n. sp.; 28, B. alta; 29, 
C. carinata; 30, C. trispinosa. 


of the protoconch is very consistent within a species, and may be used with some 
confidence to distinguish between morphologically similar species. 

Shell structure.—Two studies of the ultrastructural organization of Seguenzia 
have been published recently (Bandel 1979; Barskov et al. 1980). Bandel’s study 
investigated all aspects of shell structure from all parts of the shells of S. megalo- 
concha Rokop and S. floridana Dall (as S. monocingulata Seguenza), while Bar- 
skov et al. concentrated on the nacreous layer near the aperture of a shell of 
Seguenzia “sp. 3” from the Pacific [possibly the species cited as S. ellegans (sic) 
in Barsanova (1966)]. 

Protoconchs of S. megaloconcha and S. floridana have a very thin periostracum 
through which crystallites of the outer acicular prismatic layer protrude, forming 
nodules and irregular ridges, an organization typical of most archaeogastropods 
(except the Neritacea), but not found in mesogastropods or neogastropods (Bandel 
1979). Beneath the acicular prismatic layer are added, in order from outer to 
inner: granular, dendritic, dissected crossed acicular, and blocky prismatic layers. 
The granular and dendritic layers disappear in the post-protoconch shell, with 
the dissected crossed acicular layer forming the entire outer prismatic structure 
beneath the outer acicular layer in S. floridana; in S. megaloconcha the dissected 
crossed acicular organization is replaced by “‘spherulite sectors with marginal 
needles dissecting each other” (Bandel 1979:51). 

The nacreous layer appears only after the first two post-protoconch whorls, and 
is sandwiched between the outer dissected crossed acicular (or spherulitic) and 
inner blocky prismatic layers, except in the last whorl of actively growing indi- 


736 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


viduals where the inner blocky prismatic layer is absent. The nacreous layer 
comprises numerous lamellae of closely packed, generally rhomboidal tablets 
(Bandel 1979; Barskov et al. 1980), and forms the major structural unit of the 
adult shell (Barskov et al. 1980). Insertion of the nacre tablets into the outer 
prismatic layer may be of two types: a stair-step arrangement found in the whorl 
walls, and columnar stacks of tablets found on the columellar wall (Bandel 1979). 
Fully developed nacre, however, is the columnar nacre typical of the Pleuroto- 
mariacea and Trochacea (Bandel 1979). 

Barskov et al. (1980) reported that the shell of Seguenzia “sp. 3,’ which from 
their Fig. 1 appears very similar to S. megaloconcha, was constructed in two 
layers: 1) an outer prismatic layer with a thickness of about 0.02 mm, the structure 
of which they did not describe, and 2) an inner nacreous layer about 0.20 mm 
thick. Absence of an inner prismatic layer indicates that the specimen was not 
fully grown because the last whorl of actively growing specimens examined by 
Bandel (1979:52) also lacked the inner layer. Barskov et al. (1980) described in 
some detail the structural arrangement of the nacreous layer: generally rhomboidal 
tablets, 20-30 um long, 15—20 um wide, and 4-6 um high, closely packed into 
lamellae, with the edges of a tablet offset from those above and below it. This 
arrangement results in a stair-step, or “brickwork,” pattern of tablet stacking 
which is similar to that characteristic of some bivalves, but not previously known 
in gastropods (Barskov et al. 1980; also see Wise 1970, and Erben 1972). Barskov 
et al. interpreted this as distinguishing Seguenzia from all other known gastropods. 
However, the “brickwork” pattern described and figured by Barskov et al. appears 
virtually identical to the transitional stair-step nacre described and illustrated by 
Bandel (1979:51, pl. 3, fig. 7) which he found near the edge of the aperture. Since 
the shell chip examined by Barskov et al. was taken from near the outer lip of 
their specimen, it seems more probable that they only observed Bandel’s transition 
nacre and missed the typical gastropod type of nacre found by Bandel. Also, if 
the fracture plane is not just right, it is very difficult to distinguish between the 
different types of nacre constructions (Dr. Roger Batten, pers. comm.). Therefore, 
rather than Seguenzia being totally different in shell structure from all other 
gastropod groups, as claimed by Barskov et al., it bears strong similarity to the 
shell structures of the Pleurotomariacea and Trochacea as demonstrated by Bandel 
(1979). 

Aperture.—The aperture in Seguenzia Group I is roughly auriculate, that of 
Guttula ovate, and in the other seguenziacean genera more or less quadrate. The 
basic shape of the aperture is often distorted by relative development of, or lack 
of, a columellar tooth, and by the claw-like extension of the outer lip, especially 
in Seguenzia. Ancistrobasis is the only defined seguenziacean group which de- 
velops apertural lirae, consisting of a thickened ridge crenulated by several short, 
low, rounded ridges, and located at the abapertural edge of the posterior labral 
sinus (Fig. 20). However, several species of Seguenzia develop a ridge similar to 
that in Ancistrobasis, but without the crenulations. 


Operculum and Anatomy 


Operculum.—In all species for which the operculum has been described (e.g., 
Bayer 1971: 124, 126, Fig. 7A), it has been characterized as thin, corneous, 


VOLUME 96, NUMBER 4 737 


paucispiral with subcentral nucleus, and auriculiform. Figure 35 shows the oper- 
culum of an undescribed Seguenzia species. Superficially, it appears more similar 
to many mesogastropod opercula than to the multispiral opercula of most oper- 
culiferous archaeogastropods. However, it is perhaps unwise to attribute phylo- 
genetic significance to this structure since there are some archaeogastropods with 
similar opercula (e.g., Euchelus Philippi), and mesogastropods with multispiral 
opercula (e.g., Echininus Clench and Abbott). Moreover, the oligogyrous condition 
may be the result of the extremely small size of seguenziaceans. 

Anatomy.— Although most shell characters of apparent phylogenetic signifi- 
cance (e.g., nacreous layer, protoconch) indicate that seguenziaceans are archaeo- 
gastropod in affinity, the incompletely known anatomy of these animals shows 
an intriguing combination of characters often used to distinguish archaeogastro- 
pods from mesogastropods. There is only one published account of the anatomy 
of a seguenziacean (Guttula galatheae, Knudsen 1964); however, Dr. Philippe 
Bouchet (in litt.) has informed me that anatomical work on Carenzia carinata 
(Jeffreys) is in progress. To Knudsen’s remarks, Dr. James McLean has generously 
allowed me to add observations he made on a partial dissection of Seguenzia 
megaloconcha Rokop from the eastern Pacific, and I have added a commentary 
on my own dissection of Seguenzia sp. cf. S. eritima Verrill (hereafter S. “‘eri- 
tima’’). Knudsen’s description of the external anatomy is as follows (Knudsen 
1964:128): “The foot is broad and has a distinct median ridge. A well developed 
propodium is present. The posterior part of the foot is rounded. An epipodium 
is present, having 3—4 tentacles. The cephalic tentacles are well developed, and 
pointed. No eyes could be observed. The mantle edge has a finger-shaped tentacle 
located on the right side. The anus is situated on a rounded lobe projecting from 
the extreme right side of the mantle edge. The penis is extremely well developed.” 
McLean adds the following commentary on Seguenzia megaloconcha: ‘The foot 
is contracted so that its shape cannot be described nor the epipodial tentacles 
counted, though there seem to be six on the right and at least three, probably 6, 
on the left. In the terminology of Crisp (1981) the epipodial tentacles are papillate 
like the cephalic tentacles. The foot is too contracted to tell whether or not there 
are epipodial sense organs. Eyes and optic tentacles are lacking; the cephalic 
tentacles are long, papillate, and evenly tapered, the greatest diameter at the base 
4 times broader than the blunt tips, the attachment to the head constricted to half 
the maximum basal diameter. The right cephalic tentacle has the right subocular 
peduncle projecting from the thickened basal area. There are no neck lobes or 
cephalic lappets. The penis is longer than the cephalic tentacles, not papillate, 
evenly tapered, the tip narrower in diameter than the tips of the cephalic tentacles, 
the basal diameter exceeding the diameter of the bases of the cephalic tentacles. 
The snout is 3 the length of the cephalic tentacles, the mouth a vertical slit under 
an upper lip. The gill is attached to the mantle skirt and is pectinibranch with 
about 30 triangular filaments, with no evidence of a free bipectinate tip or the 
transverse pallial vein of the Trochacea. The edge of the mantle skirt is finely 
fringed, but the pallial tentacle on the right side of the mantle skirt is relatively 
small.” 

The following description is based on my study of a male specimen of S. 
“eritima’’: Although strongly contracted, the foot appears to be truncated ante- 
riorly, tapering gradually to a broadly rounded, possibly bilobed, posterior; there 


738 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Figs. 31-34. Animal of Seguenzia “eritima’’: 31, animal removed from shell, right side; 32, same, 
left side; 33, mantle reflected to right showing head and anterior portion of pallial cavity; 34, dorsal 
view of intestine and kidney. Scale bar = 1 mm for Figs. 31, 32; 1.5 mm for Figs. 33, 34. Abbreviations: 
a, anus; ap, accessory cephalic process; as, anal sinus; bs, basal sinus; cl, lateral processes of snout; 
cm, columellar muscle; cs, columellar sinus; ct, ctenidium; dg, digestive gland; et, epipodial tentacle; 
ev, efferent branchial vein; f, foot; i, intestine; k, kidney; It, left cephalic tentacle; m, mouth; op, 
operculum; p, penis; r, rectum; rt, right cephalic tentacle; sop, subocular peduncle; st, stomach; t, 
testis. 


is a small, short tentacle at each anterolateral corner. There appear to be four 
pairs of epipodial tentacles which increase in size from anterior to posterior. 
Epipodial sense organs appear to be lacking. Epipodial tentacles (Fig. 32, et) are 
like those described by Crisp (1981). The operculum is very thin, corneous, with 
a subcentral nucleus and about four or five rapidly expanding whorls. 

The snout is very short, but provided with long, triangular, lateral extensions 
(oral lappets?; Figs. 32, 33, cl). The mouth is transversely elongate, surrounded 
by an outer lip which is interrupted mid-ventrally. The cephalic tentacles (Figs. 
31-33, rt, It) are long, papillate, and tapered from a broad base to a blunt tip. 
The right tentacle bears a prominent subocular peduncle (Figs. 31, 33, sop) on 


VOLUME 96, NUMBER 4 739 


the base. There are no eyes or optic tentacles. A very long, slender penis (Figs. 
31, 33, p) arises from the right side of the head lateral and posterior to the right 
cephalic tentacle; a ciliated sperm groove runs along the ventral side of the free 
part of the penis, becoming lateral at the base where the seminal duct opens into 
the groove. Two small accessory tentacles are situated just posterior to the base 
of the penis. A large cephalic process (Figs. 31, 33, ap) arises dorsal and posterior 
to the left cephalic tentacle, crosses the head obliquely to the right, and curves 
forward to the right of the right cephalic tentacle. It is attached to the head for 
about half its length, becomes free behind the right tentacle and projects forward 
between the tentacle and penis. 

The mantle edge bears two deep and two shallow papillate embayments, or 
mantle sinuses, which correspond in position to the shell sinuses. One of the major 
embayments, corresponding to the basal sinus of the shell, is located over the 
base of the left cephalic tentacle and undoubtedly acts as the primary incurrent 
siphon (Fig. 32, bs). This embayment is papillate along the entire edge, and a 
ridge of muscle is located just back of the edge, indicating that this area can be 
expanded to some extent. The other large embayment functions as the excurrent, 
or anal, siphon, and is in the extreme right part of the mantle edge (Figs. 31, 33, 
as). It too is papillate, but only along the posteriormost edge. The medial edge of 
this mantle sinus is strongly folded, indicating that considerable extension is 
possible, forming a baffle or pseudosiphon directing the exhalent current to the 
right. The two smaller embayments, one located on the extreme left (Fig. 32, cs) 
and the other mid-dorsally, are both papillate; the other areas of the mantle edge 
are smooth. There are no pallial tentacles. 

The ctenidium (Figs. 32, 33, ct) is monopectinate, with about 30 triangular 
lamellae, extending in a curve from a posterolateral position medially and ante- 
riorly to end just to the right of the large left mantle sinus. Neither osphradium 
nor hypobranchial gland was observed. The intestine (Figs. 3 1-34, 1) is prominent, 
convoluted, arising from the style sac of the stomach, running forward to about 
the accessory cephalic process, turning to the left and running back to the middle 
part of the stomach, and finally turning forward along the right to the anus which 
is On a small papilla. The intestine was filled with fine detrital material (mostly 
mud, foraminiferan tests, and diatom frustules) which was consolidated into a 
continuous fecal string which had an ovoidal cross section and a generally dorsal 
longitudinal groove. The preservation of the large, thin-walled stomach (Figs. 31, 
32, st) was not adequate for detailed study of the internal morphology. The left 
kidney (Fig. 34, k) is located above the stomach, above and within the posterior 
bight of the intestine. There appears to be no right kidney. The circulatory system 
is apparently of monotocardian layout. The heart is located dorsolaterally to the 
right of the anterior end of the stomach. The single auricle receives the long 
efferent branchial vein and the ventricle is not penetrated by the intestine. The 
aortae emerge and run along the right side of the body. 

The above descriptions, although of differing completeness, have several ele- 
ments in common. The epipodium, epipodial tentacles, and penis are shared by 
all three species. A pallial tentacle is present in both Guttula galatheae and Se- 
guenzia megaloconcha. Although by no means conclusive, these similarities, along 
with radula and shell characters, tend to support my inclusion of Guttul/a in the 
Seguenziacea. 


740 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


The descriptions of Seguenzia megaloconcha and S. “‘eritima”’ agree in most 
respects, but there are a few obvious differences in external morphology between 
the two species. The pallial tentacle of S. megaloconcha is absent in SS. “‘eritima,” 
and there is no basal constriction of the cephalic tentacles in S. “‘eritima.” Presence 
of pallial tentacles in one and not another species of the same genus is also seen 
in the rissoacean genera Cingula and Onoba (Fretter and Graham 1978b). This 
situation may also be found in individuals of the same species (e.g., Onoba semi- 
costata; Fretter and Graham 1978b:165). The basal constriction of the cephalic 
tentacles in S. megaloconcha may not have any significance but bears investigation 
in other species. The lateral extensions of the snout in S. “eritima,” not found in 
S. megaloconcha, are of functional importance but are of unknown taxonomic 
significance at present. The extensions are probably primarily sensory in nature, 
supplementing the cephalic tentacles, but may be used in feeding, either by stirring 
up the substrate, or by aiding in the movement of food into the mouth. 

The most striking difference between S. megaloconcha and S. “‘eritima’’ is the 
large accessory cephalic process which lies obliquely across the head of S. “‘eri- 
tima,”’ but is totally absent in S. megaloconcha. It is a conspicuous structure, 
arising from the head behind the left tentacle, remaining attached to the dorsum 
behind the right tentacle, and finally projecting forward as a finger-like process 
to the right of the right tentacle. The internal structure of this process has not yet 
been determined, although it appears to be solid and muscular; its function is not 
immediately evident, and I have not been able to locate any reference in the 
literature to a similar structure occurring in any other prosobranch. The penes of 
S. megaloconcha and S. “eritima’’ originate at different positions. That of S. 
megaloconcha arises from the dorsum of the head near the midline, and that of 
S. “eritima’ arises from the right side. The significance of this difference is unclear 
to me. The location of the anterior and posterior aortae, on the right side of the 
animal, is apparently unique among coiled gastropods. All accounts I have been 
able to locate have described the aortae being on the left side. Knowledge of the 
anatomy of other seguenziacean species is needed before the systematic signifi- 
cance of these anatomical structures can be inferred. 


Radula 


The distinctive seguenziacean radula combines features of rhipidoglossate and 
taenioglossate radular types, but fits neither type. All seguenziaceans for which 
radulae have been illustrated (Figs. 39-48) have the central part of the radula 
comprised of a rhachidian (Figs. 37, 38, rh) flanked by a single pair of laterals 
(Figs. 37, 38, 1), and the outer part of 4-12 pairs of marginals which completely 
cover the central portion in the folded, non-working condition (Fig. 36). It thus 
appears to be a modification of the rhipidoglossate radula, derived by reduction 
of number of laterals and marginals to the point where, in Seguenzia Group I, 
the radula bears a strong superficial similarity to the taenioglossate state (Fig. 46). 
In fact, many of the species in the family Triphoridae have radular formulae 
(Triphorinae, 30-9.1.1.1.9-30; Mastoniinae, 8-5.1.1.1.5-8; see Kosuge 1966) which 
are similar to that of the seguenziaceans (12-4.1.1.1.4-12). However, the teeth of 
the two radular types are totally different structurally, and there is little resem- 
blance in shell or anatomy between triphorids and seguenziaceans. There is little 


VOLUME 96, NUMBER 4 741 


Figs. 35-38. SEM micrographs of operculum and radula of Seguenzia sp.: 35, operculum, 47 x: 
36, intact radular ribbon with marginals folded over central part, 268 x: 37, individual teeth teased 
out of ribbon, 570 x; 38, rhachidian and lateral teeth, 1340. 


doubt that these two groups are unrelated, and further comparisons are unnec- 
essary. There are a large number of modified radular types in the archaeogastro- 
pods, especially in deep-water groups, and use of this structure to infer relation- 
ships should be made with care. 

Seguenzia.—(Figs. 36-38, 46-48). The Seguenzia radula consists of the rhach- 
idian, a single pair of laterals, and (as far as is known) four pairs of marginals per 
row. The rhachidian is pyriform with a basal attachment process and a single, 
denticulate cusp. The lateral has a broad, rather triangular base with a long, slender, 
finely denticulate cusp near the proximal corner. The first marginal is larger than 
the rest, blade-like, and denticulate on both sides near the tip. The remaining 
marginals are long, very slender, with a few fine, spinular teeth slightly removed 
from the tip, and a rhomboidal base. Both Schepman (1909) and Barnard (1963c) 
illustrated Seguenzia radulae (S. melvilli and S. simplex; Figs. 47 and 48, re- 
spectively) showing rhachidians without the basal process and cuspless laterals. 
In S. melvilli, Schepman may have simply missed the basal process of the rhach- 
idian, and the cusp of the lateral may have been worn or broken off. Barnard’s 
species, S. simplex, may not be congeneric with Seguenzia sensu stricto, although 
the same considerations as those mentioned for S. me/villi may also apply here. 
Scanning electron micrographs presented by Bandel (1979) and Hickman (1980) 
of the radula of S. megaloconcha are almost indistinguishable from those pre- 


742 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


sented here (Figs. 36-38, 46) of the radula of an undescribed species from the 
Philippines. Schepman (1908) also described the radula from a syntype of Basilissa 
lampra Watson, which he mistakenly considered the type of Basilissa. However, 
that radula (Fig. 45) is more similar to those described for Seguenzia (Figs. 46- 
48) than to those of true Basilissa (Figs. 40, 41). The shell shape of B. Jampra is 
similar to those of Seguenzia Group III, so it is probable that this species is a 
Seguenzia sensu lato rather than a Basilissa. Thiele (1925) described the radula 
of Fluxina trochiformis Schepman, in which he found a single, broad lateral, a 
single, rather wide inner marginal, and five outer marginals denticulate on the 
distal edges. Thiele therefore transferred F. trochiformis to Basilissa. This species 
bears a strong conchological resemblance to F. discula Dall, which has recently 
been assigned to Basilissa (Merrill 1970b; Quinn 1979). However, F. discula and 
F. trochiformis are here referred to Seguenzia Group IV pending description of 
that group as a new genus by Marshall (in press). 

Basilissa. — (Figs. 40, 41). The radula of Basilissa differs from that of Seguenzia 
sensu stricto in having 6-7 marginal teeth (Bayer 1971) and a larger, more tri- 
angular cusp on the lateral tooth. 

Ancistrobasis.—(Fig. 39). The radula of Ancistrobasis has not been previously 
illustrated or described. It is more trochoid than that of either Seguenzia or 
Basilissa, having a rather large, rectangular rhachidian with denticulate cusp, a 
large, subquadrate lateral with an inwardly directed cusp denticulate on both sides, 
and twelve slender marginals finely denticulate near the tip. 

Guttula. —(Figs. 42—44). The radulae of all three nominal species of Guttula 
have been described and illustrated, and indicate an affinity with more typical 
seguenziaceans. The rhachidian is broad, rather rectangular, with a denticulate 
cusp. The lateral is large, broad, and rectangular, with or without a cusp. There 
are a “few” marginals (no one has given a specific number, although Barnard 
(1963c:266) states “not more than ten’’). The length of the radula of G. sibogae 
Schepman is about 1 mm and its width about 0.3 mm (Schepman 1908). Schep- 
man counted about 20 rows of teeth, Barnard (1963c) about 22. 


Fossil Record 


As far as is known, the Seguenziacea are relatively young, possibly originating 
sometime during the late Cretaceous or early Tertiary, and at least one group 
(Ancistrobasis) attained its modern form by the Eocene. Over 20 fossil species 
have been described, most of which were assigned to Basilissa, but less than half 
of these taxa are true seguenziaceans (Tables 1, 2). 

The earliest occurrences of the Seguenziacea in the geological record are Se- 
guenzia radialis Tate (Tate 1888, 1890) and Basilissa cossmanni Tate (Tate 1894) 
from the Eocene of South Australia, and Basilissa (Ancistrobasis) pacifica Ladd 
(Ladd 1970) from the Eocene of Tonga. All three taxa are here considered An- 
cistrobasis species. Ancistrobasis also occurs in the Pliocene of Sicily (Solarium 
reticulatum; Philippi 1844; Jeffreys 1885). The first known occurrence of Se- 
guenzia is S. hapala Woodring from the middle Miocene of southern Mexico 
(Perrilliat 1972). Specimens of this species have also been collected from the basal 
Pliocene of the Dominican Republic (Gurabo Formation; H. E. Vokes, in Iitt.), 
the upper Pliocene-basal Pleistocene of Jamaica (Bowden Formation; Woodring 


VOLUME 96, NUMBER 4 743 


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/ Me \\ oS Ze | is 
Wee. 
39 Wy ON . He J ee 
mo) iL Lo fr AN \ \ 
oi (esc) ~ \ 
40 of V \ lt carta ig em) AW 
y dale \ ‘ Se 
At a) ae (1 
pO j | | 
ae a crime, 


Figs. 39-48. Diagramatic sketches of seguenziacean radulae, not to scale: 39, Ancistrobasis de- 
pressa;, 40, Basilissa alta (after Bayer 1971); 41, B. sibogae (after Schepman 1908); 42, Guttula sibogae 
(after Schepman 1908); 43, G. blanda (after Barnard 1963); 44, G. galatheae (after Knudsen 1964); 
45, Seguenzia lampra (after Schepman 1908); 46, Seguenzia sp.; 47, S. melvilli (after Schepman 1909): 
48, S. simplex (after Barnard 1963). 


1928), and the Recent throughout the Gulf of Mexico and Caribbean [Treece, 
1977, 1980 (as S. formosa); Treece 1979 (as S. monocingulata); Quinn unpub- 
lished data]. Seguenza (1876, 1877, 1879) recorded S. monocingulata from the 
Pliocene of Italy. Mioseguenzia Nordsieck, 1973, described with a Miocene type- 
species, 1s cypraeacean (Bouchet, in litt.), not seguenziacean. Some members of 
the Paleozoic-Mesozoic family Omphalotrochidae Knight, 1945, bear a superficial 
resemblance to Basilissa, but, as discussed below, this similarity is probably 
convergent and not an indication of relationship. 

Other allocations of fossil species to Basilissa (Cossmann 1888; Noetling 1901; 
Oppenheim 1906, 1923; Yokoyama 1922; Koperberg 1931; Riedel 1932; Ravn 
1933, 1939) followed Cossmann’s (1888) erroneous concept of the genus. Most 
of these species appear to be referrable to the Trochidae (Table 2). 


744 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Table 2.—Taxa rejected from, or of doubtful inclusion in, the Seguenziacea. 


ORIGINAL BINOMEN 


Seguenzia tricarinata Jeffreys, 1885 Nassariid protoconch 

S. laxa Jeffreys, 1885 Indeterminate 
+Basilissa boutillieri Cossmann, 1888 2% 

B. patula von Martens, 1901 Calliotropis 
+B. lorioliana Noetling, 1901 ? 

B. ottoi var. chuni von Martens, 1903 Calliotropis 
+Trochus (Basilissa) libycus Oppenheim, 1906 2 

B. babelica Dall, 1907 Orectospira 

B. niceterium Hedley and May, 1980 Calliostoma 
+B. ?laeviuscula Yokoyama, 1922 Proconulus? 

Trochus (Basilissa) lukavatzensis Oppenheim, 1923 % 
+B. lemoinei Koperberg, 1931 ? 


+B. antiqua Riedel, 1932 
+B. tricincta Ravn, 1933 
+B. odumi Ravn, 1939 


+B. (Orectospira) nenokamiensis Kanno, 1958 


B. bicarinata Habe, 1961 


9 
ef. Bathybembix 
cf. Bathybembix 
Orectospira 
Calliotropis 


Mioseguenzia Nordsieck, 1973 
Type-species. —Janthina cimbrica Sorgenfrei, 1958; by original designation. 


+M. cimbrica (Sorgenfrei, 1958) Cypraeacean larval 
shell 

M. cimbrica recens Nordsieck, 1973 Cypraeacean larval 
shell 

M. conica Nordsieck, 1973 Cypraeacean larval 
shell 


+ = species described from fossil material. 
? = affinities uncertain. 


Relationships 


Speculations on relationships of the Seguenziacea to other prosobranch groups 
have been, and remain, inconclusive. Jeffreys (1876, 1879) strongly advocated 
assignment of Seguenzia to the Solariidae (=Architectonicidae), but this view has 
never gained acceptance. Following Seguenza’s (1876) lead, many authors have 
placed Seguenzia in the Trochidae (Watson 1879a, 1886; Thiele 1925, 1929; 
Wenz 1938; Cotton 1959; Keen and Cox 1960; Barnard 1963c; Knudsen 1964; 
Barsanova 1966; Bayer 1971; Abbott 1974). These authors also considered Bas- 
ilissa and Guttula to be trochids. Seguenzia has also been included in the Pleu- 
rotomariidae (Tryon 1883), Haliotidae (von Martens 1881; Tate 1888), and Scis- 
surellidae (Locard 1898), principally because of the posterior sinus in the outer 
lip. Verrill’s (1884) establishment of the Seguenziidae, including Seguenzia and 
Basilissa, and placement in the Mesogastropoda near the genus Aporrhais was 
followed by Golikov and Starobogatov (1975). The Seguenziidae have also been 
placed near, or included in, the Trichotropidae (Tryon 1887; Dall 1889b, c; Tate 
1890; provisionally by Schepman 1909), near the Triphoridae (Dall 1927; Wood- 
ring 1928; Taylor and Sohl 1962), between the Archaeogastropoda and Meso- 
gastropoda (Keen 1971), and near the Trochidae (Clarke 1961, 1962; Bandel 1979; 
Quinn 1979, 1981; Boss 1982). Recently Goryachev (1979) proposed inclusion 


VOLUME 96, NUMBER 4 


745 


Table 3.—Comparison of shell and anatomical features of the Trochacea, Seguenziacea, and Ris- 
soacea. Characters of the Trochacea and Rissoacea taken from Fretter and Graham (1962). 


Structure 


Trochacea 


Seguenziacea 


Rissoacea 


Shell Nacreous or porcella- Nacreous Porcellaneous 
neous 
Protoconch Archaeogastropod Archaeogastropod Mesogastropod 
Radula Rhipidoglossate Modified rhipidoglos- Taenioglossate 
sate 
Epipodium Present, with tentacles Present, with tentacles Absent 
and sense organs 
Reproductive No copulatory organs; Penis and closed gon- Penis present; glandular 
system gametes shed oduct present; fer- gonoducts with acces- 
through right kid- tilization internal sory structures in fe- 
ney; fertilization ex- male; fertilization in- 
ternal ternal 
Ctenidium Bipectinate (rarely Monopectinate Monopectinate 
monopectinate) 
Pallial Absent Present in at least 2 Often present 
tentacles species 
Circulatory Diotocardian, with Monotocardian (?), Monotocardian 
system transverse pallial without transverse 
vein pallial vein 
Kidneys Both right and left Right kidney lost (?) Right kidney lost 
present 
Intestine Long, with anterior Long, with anterior Short, without anterior 


loop; rectum pass- 
ing through ventri- 


loop; rectum free of 
ventricle 


loop; rectum free of 
ventricle 


cle 


of the Seguenziidae in the Protopoda Fisher, 1884 [sensu Golikov and Starobo- 
gatov (1975) = Turritellacea + Vermetacea] or possibly in a separate order. In 
view of data presented here, both proposals seem very unlikely. These assignments 
were made principally on the evidence of shell and, occasionally, radular char- 
acters. Therefore, features of the anatomy and shell morphology detailed in this 
paper permit a more critical examination of the possible affinities of the Seguen- 
ziacea than has yet been possible. 

The first problem in assessing the probable relationships of the Seguenziacea 
is to determine the proper prosobranch suborder, Archaeogastropoda or Meso- 
gastropoda, to which the group should be assigned. Table 3 presents a summary 
of the known characters of the Seguenziacea in comparison with corresponding 
traits of the Trochacea and Rissoacea, as representatives of the Archaeogastropoda 
and Mesogastropoda, respectively. If the use of the term “mesogastropod”’ is 
restricted to describe a certain level of anatomical organization rather than as a 
discrete taxonomic entity (Fretter et al. 1981), the Seguenziacea must be inter- 
preted as mesogastropodan. However, as will be argued below, most of the Se- 
guenzia-mesogastropod (the taxon) similarities are probably convergent, and may 
not be indicative of any phytogenetic relationship. Therefore, I propose that the 
Seguenziacea are highly modified and specialized archaeogastropods whose an- 


746 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


atomical advances are consequences of extremely small body size and adaptation 
to life in the deep-sea. 

Two seguenziacean characters which are the most indicative of archaeogastro- 
pod affinity are the shell and radula. Nacreous shells are known in the Monopla- 
cophora and some lower groups of the Bivalvia, Cephalopoda, and Gastropoda, 
leading to the assumption that nacre is a primitive character within the Mollusca. 
Besides the Seguenziacea only two groups of living gastropods, the Pleurotomar- 
iacea and most Trochacea (both archaeogastropod), have nacreous shells. Nacre 
has not been noted in any mesogastropod or higher group. The seguenziacean 
radula seems to be a modification of the rhipidoglossate radula. Verrill’s (1884) 
description of the radula of S. eritima as taenioglossate has induced many authors 
to assign Seguenzia to the Mesogastropoda. The seguenziacean radula does re- 
semble the taenioglossate radula superficially, principally in having only a single 
lateral and a reduced number of marginals in each radula half-row. Of the meso- 
gastropod families discussed by Boss (1982), only two were recorded as having 
more than two marginal teeth, the Turritellidae (3-0.1.1.1.0-3) and the Triphor- 
idae (30-9.1.1.1.9-30 or 8-5.1.1.1.5-8), and neither family is in any way similar 
to seguenziaceans in shell or anatomy. Moreover, the two marginals of the typical 
taenioglossate radula are very similar to each other in shape and size. Seguenzia- 
cean marginals are morphologically different, the innermost tooth being rather 
large and robust and the outer whisker-like, a situation reminiscent of several 
rhipidoglossate ground plans in which the marginals may be divided into two or 
more distinct morphological groups, with the innermost usually much stronger 
than the outer. It has been suggested to me that the multiple marginals of the 
seguenziacean radula might have been produced by repeated splitting of the tae- 
nioglossan marginals. While this remains a possibility, especially in view of the 
observations of Shimek and Kohn (1981) on the Turridae, the apparent affinity 
of the Seguenziacea with archaeogastropods such as the Trochacea (discussed 
below) suggests that the seguenziacean radula is merely a modification of the 
rhipidoglossate radula, and is adapted for sweeping particulate matter from the 
bottom and conveying it to the esophagus. 

Other traits of the Seguenziacea which suggest archaeogastropod affinity are a 
long intestine with an anterior loop, a subocular peduncle on the base of the right 
cephalic tentacle, and an epipodium with epipodial tentacles. The intestine follows 
a course similar to that of the Trochacea, but the seguenziacean anterior loop 
(perhaps not homologous with that of the Trochacea?) is much longer, and the 
intestine does not penetrate the ventricle. The subocular peduncle has been noted 
in several species of Trochidae (Crisp 1981), but to my knowledge is not known 
in any other prosobranch group, except, perhaps, the Fissurellacea (McLean, in 
litt.). The epipodium and associated tentacles have been generally considered an 
archaeogastropod trait (Fretter and Graham 1962), but epipodial tentacles are 
also known in some mesogastropods [e.g., Litiopidae and Dialidae (Houbrick 
1980; see also Thiele 1929)]. Even the fecal string of Seguenzia resembles that of 
the Trochacea, although there is no liver string (see Fretter and Graham 1962). 

Several seguenziacean features show progression into an advanced grade of 
organization. However, most of the advancement may be attributed to adaptation 
to a small body size and for increased functional efficiency. The radula has already 
been discussed. The other principal modifications include the development of an 


VOLUME 96, NUMBER 4 747 


advanced reproductive tract, a monopectinate ctenidium, and functional inhalent 
and exhalent pallial siphons. 

Although no in-depth anatomical examination of the seguenziacean reproduc- 
tive system has yet been made, presence of a well-developed penis suggests other 
modifications of the tract, such as possible accessory structures in the female 
oviduct. I have not examined any females, but Dr. Anders Warén (pers. comm.) 
has indicated that such modifications may be present in female Carenzia carinata 
from the northeastern Atlantic. My examination of the male of S. “‘eritima’’ 
indicates a simple sperm duct, without prostate gland, which opens into a ciliated 
sperm groove on the penis. Penial structures, either as modifications of a cephalic 
tentacle or de novo structures, are rare, but not unknown in the archaeogastropods. 
Perhaps the best known situation is in the Neritacea, in which a true cephalic 
penis has been developed (see Fretter and Graham 1962; Fretter 1965). Several 
other archaeogastropod groups also have been reported to have a penis: Coccu- 
linacea (Cocculina and Addisonia; see Dall 1889b, 1890); Trochidae [Solariella 
(=Calliotropis) and Turcicula (=Bathybembix),; Dall 1889a, b, 1890]; Fissurellacea 
(Rimula and Fissurella; Dall 1889b); and the recently described Neomphalacea 
(Neomphalus,; McLean 1981; Fretter et al. 1981). The Cocculinacea and Neom- 
phalacea each have a modified cephalic tentacle (the former the right, and the 
latter the left tentacle), but the function as a penis is undoubted. The “‘penis”’ 
described by Dall (1889b) in the Fissurellacea was found to be solid by Odhner 
(1932) who referred to the structure as a “‘sexual cirrus.”’ The rudimentary “penis” 
reported in the Trochidae (Dall 1889a, b, 1890) has not been investigated sub- 
sequently. McLean (in litt.) considers the penis-like structures of the fissurellaceans 
and trochids to be the right subocular peduncle. Since all archaeogastropod groups 
with an undoubted penis except the Neritacea (Cocculinacea, Neomphalacea, and 
Seguenziacea) are principally deep-sea forms, and most are extremely small (less 
than 10 mm, except Neomphalus fretterae McLean, 1981, which may be as much 
as 30 mm in diameter and which lives in a unique situation, the Galapagos Rift), 
energy conservation is a primary concern. Internal fertilization severely reduces 
the wastage of gametic products, and evolution of copulatory structures would be 
the most efficient solution to that problem. 

A monopectinate ctenidium is generally associated with a mesogastropod, or 
higher, grade of organization, and has not been reported previously in any ar- 
chaeogastropod except Umbonium (Fretter 1975; McLean 1981). Presence of such 
a ctenidium in the Seguenziacea may well be an adaptation for economy of space 
in the pallial cavity of these small animals. A monopectinate ctenidium provides 
more efficient use of space and eliminates any circulatory dead spots which are 
found in most bipectinate ctenidia. However, the change from bipectinate to 
monopectinate conditions must require some compensation for the decrease in 
strength of the ciliary currents flowing through the pallial cavity caused by re- 
duction of the number of ctenidial leaflets; therefore, the Seguenziacea, like many 
mesogastropods and virtually all neogastropods, have modified the mantle edge 
to form inhalent and exhalent siphons. However, seguenziacean siphons are pa- 
pillate embayments in, rather than extensions of, the mantle edge. The papillae 
may augment the ctenidial cilia in producing the inhalent and exhalent currents, 
and the siphons undoubtedly enhance the efficiency and directionality of the 
currents. The seguenziacean siphons are apparently analogous to those of meso- 


748 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


gastropods and neogastropods, but are de novo structures derived in conjunction 
with evolution of a monopectinate ctenidium. In addition, in at least one species 
(S. “‘eritima’’), a cephalic process forms a baffle across the opening of the mantle 
cavity which probably prevents water from entering or exiting the mantle cavity 
except through the siphons. 

Since the known features of the Seguenziacea are either clearly archaeogastropod 
in nature or can be derived from archaeogastropod structures as adaptations for 
small body size and life in the deep-sea, it seems most prudent to consider the 
Seguenziacea the end of a long-separated evolutionary lineage of archaeogastropod 
derivation, which has developed mesogastropod-like adaptations independent of 
any mesogastropod lineage. In this regard, the Seguenziacea are similar to the 
Neomphalacea (McLean 1981) and Neritacea (Fretter and Graham 1962; Fretter 
1965). 

If Seguenziacea are indeed archaeogastropods, as argued here, then which group 
might be considered seguenziacean precursors? Solely on the basis of shell shapes 
and position of anal sinuses, two possibilities exist. First is an origin in a euom- 
phalacean group such as the Omphalotrochidae Knight, 1945, especially Om- 
phalotrochus Knight, 1945, Orecopia Knight, 1945, and Babylonites Yochelson, 
1956. Earlier (Quinn 1981) I suggested that the Seguenziacea might have been 
derived from such omphalotrochids based on the subsutural sinus and channeled, 
claw-like extension of the outer lip, which are apparent homologues of the anal 
and basal sinuses of the seguenziacean shell. McLean (1981) has since pointed 
out that the Omphalotrochidae are not known to have nacreous shells, and that 
a reversion to the nacreous condition would be unlikely. In addition, there is a 
gap of about 200 million years between the extinction of the Omphalotrochidae 
(Middle Permian) and the earliest known seguenziacean (Eocene). Therefore, an 
Omphalotrochidae-Seguenziacea lineage is not supported. 

The other, and more probable origin, involves a derivation from within the 
Pleurotomariacea- Trochonematacea-Trochacea lineage. It has. been hypothesized 
that the Pleurotomariacea gave rise to the Trochonematacea (Knight et al. 1960), 
which in turn were ancestral to the Trochacea (Fretter and Graham 1962). All 
three superfamilies have nacreous shells and the Pleurotomariacea and Trocho- 
nematacea both have labral sinuses. We know the anatomy of living Pleuroto- 
mariacea and Trochacea from which we may infer possible relationship with the 
Seguenziacea, but inferences about the extinct Trochonematacea are purely spec- 
ulative. The Pleurotomariacea are gastropods retaining such primitive conditions 
as paired pallial organs, simple reproductive system and intestine penetrating the 
ventricle [see Fretter and Graham (1962) and Fretter (1964, 1966) for summary 
of other features]. From the position of the shallow anal sinus or channel, the 
Trochonematacea were probably dibranchiate, with the right ctenidium possibly 
being lost in the later forms (Knight et al. 1960). The Trochacea have lost the 
right ctenidium and osphradium, but retain the other paired pallial structures, 
the intestine is long, with an anterior loop, but still penetrates the ventricle, and 
the reproductive system remains relatively unspecialized (Fretter and Graham 
1962). The Pleurotomariacea and Trochacea have rhipidoglossate radulae, al- 
though comparatively reduced in the latter group, and, in most other anatomical 
regards, the Trochacea and Pleurotomariacea are also remarkably similar (Fretter 
and Graham 1962; Fretter 1964, 1966; Graham 1965). Although similar to the 


VOLUME 96, NUMBER 4 749 


Trochacea and Pleurotomariacea in having nacreous shells, rhipidoglossate rad- 
ulae (although modified), epipodium and epipodial tentacles, and an anterior loop 
of the intestine, the Seguenziacea have highly modified reproductive and circu- 
latory systems and a monopectinate ctenidium, which indicate a long-standing 
separation from the mainstream of archaeogastropod evolution. My contention 
that the seguenziaceans are more likely to have had their origin within the Tro- 
chacea rests primarily on the fact that it seems more plausible to derive the 
seguenziacean organization as modifications of a trochoid organization than of a 
zeugobranch condition, and secondarily on the radula, trochoid-like fecal-string, 
and cephalic tentacles, especially the presence of a subocular peduncle on the right 
tentacle. Admittedly, this is rather tenuous evidence from which to draw a con- 
clusion, but may serve as a working hypothesis for further research. 

Since the anatomy of no seguenziacean is fully known, only three having been 
partly described, and the fossil record is so scanty, no discussion of relationships 
within the superfamily is now possible, nor is it possible to determine whether 
the anal sinus is a primitive character which was inherited from some ancestor 
or an innovation of the seguenziaceans. In view of the tendency of many other 
prosobranchs to develop similar structures, I believe the latter to be true, thus 
reinforcing my belief that the seguenziaceans are derived from some holosto- 
matous ancestor, 1.e., a trochoid. It is probable that additional family-group taxa 
may be required as future systematic studies are completed. Indeed, the mono- 
phyly of the Seguenziacea may even now be questioned. For example, it could 
be argued that the genera with labral sinuses may have evolved from the Tro- 
chonematacea, Guttula may have arisen independently from the Trochacea, and 
the selective pressures of similar habitats subsequently produced similarly con- 
structed organisms. However, the presence of a penis, epipodium and epipodial 
tentacles, and radulae and shells similar to some undoubted seguenziaceans, sug- 
gest that Guttula should be included in the Seguenziacea. 

Therefore, the Seguenziacea, as here defined, comprise the genera Seguenzia, 
Carenzia, Basilissa, Basilissopsis, Ancistrobasis, Thelyssa, and Guttula. The su- 
perfamily is characterized by nacreous shells, generally with labral sinuses, mod- 
ified rhipidoglossate radula, and an anatomy retaining some archaeogastropod 
traits (epipodium with tentacles, long anterior loop of the intestine, and subocular 
peduncle) but attaining several mesogastropod features (penis in males, mono- 
pectinate ctenidium, and monotocardian circulatory system). This combination 
is so different from other known prosobranchs that there can be little doubt that 
recognition of a separate superfamily as suggested by Keen (1971), Golikov and 
Starobogatov (1975), Goryachev (1979), Quinn (1981, 1983a, b), McLean (1981), 
and Marshall (in press) is necessary. The Seguenziacea are here retained within 
the Archaeogastropoda as an independent offshoot of the Pleurotomariacea-Tro- 
chonematacea-Trochacea lineage, with the Trochacea as the most probable an- 
cestral stock. 


Acknowledgments 


I thank the following curators for the opportunity to examine specimens in 
their care: Dr. Richard S. Houbrick (U.S. National Museum of Natural History, 
Mollusks), Dr. Gilbert L. Voss (Rosenstiel School of Marine and Atmospheric 
Science, University of Miami), Dr. Kenneth J. Boss (Museum of Comparative 


750 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Zoology, Harvard University), Dr. John Peake and Ms. Kathie Way (British 
Museum, Natural History), Dr. Thomas R. Waller (U.S. National Museum of 
Natural History, Paleobiology), Mr. Granville D. Treece (Marine Science Institute, 
University of Texas), and Dr. Linda H. Pequegnat (Texas A&M University). I 
am especially indebted to Dr. Michael A. Rex (University of Massachusetts) for 
providing the preserved specimens of Seguenzia “‘eritima.” 

Parts of this paper were presented at the 1980 and 1982 annual meetings of 
the American Malacological Union. I am grateful to those colleagues who provided 
numerous valuable critiques of and suggestions on those presentations. Special 
thanks go to Drs. James H. McLean (Los Angeles County Museum), Carole S. 
Hickman (University of California, Berkeley), Philippe Bouchet (Laboratoire de 
Malacologie, Muséum National d’Histoire Naturelle, Paris), and Boss and Houb- 
rick who kindly read and commented on earlier versions of this manuscript. Dr. 
McLean also generously allowed me to incorporate some of his unpublished 
anatomical observations and examine specimens of Seguenzia megaloconcha. Dr. 
Bruce A. Marshall (National Museum of New Zealand) kindly provided me with 
advance information concerning new taxa from the New Zealand region. The 
manuscript also benefitted from comments by William G. Lyons, Thomas H. 
Perkins, and Scott A. Willis, and SEM services were provided by Lana Tester (all 
Florida Department of Natural Resources Bureau of Marine Research). 

This paper is a scientific contribution from the Rosenstiel School of Marine 
and Atmospheric Science, University of Miami, and forms part of a dissertation 
submitted to the school in partial fulfillment of the requirements for the degree 
of Doctor of Philosophy. 


Literature Cited and Bibliography 


Abbott, R. T. 1974. American seashells. 2nd Ed. Van Nostrand Reinhold, New York, 663 pp. 

Bandel, K. 1979. The nacreous layer in the shells of the gastropod-family Seguenziidae and its 
taxonomic significance. — Biomineralisation 10:49-61. 

Barnard, K. H. 1962. New species and records of South African marine Mollusca from Natal, 

Zululand, and Mogambique.— Annals of the Natal Museum 15(19):247-254. 

. 1963a. Deep-sea Mollusca from the region south of Madagascar. — Department of Commerce 

and Industries, Investigational Report No. 44:1-19. 

—. 1963b. Deep-sea Mollusca from west of Cape Point, South Africa.—Annals of the South 

African Museum 46:407-453. 

. 1963c. Contributions to the knowledge of South African marine Mollusca. Part IV. Gastropo- 

da: Prosobranchiata: Rhipidoglossa, Docoglossa, Tectibranchiata, Polyplacophora, Soleno- 

gastres, Scaphopoda.— Annals of the South African Museum 47:201-360. 

. 1966. Deep-sea Mollusca from the region south of Madagascar. — International Indian Ocean 

Expedition, Collected Reprints 2:1—19 (reprint of Barnard, 1963a). 

. 1974. Contributions to the knowledge of South African Marine Mollusca. Part VII. Revised 

Fauna List.—Annals of the South African Museum 47:663-781. 

Barsanova, N. G. 1966. K nachozhdeniyu glubokovodnikh predstavitelei semeistva Seguenziidae 
(Gastropoda, Prosobranchia) v Tikhom Okeane. [On the finding of deep-sea representatives of 
the family Seguenziidae (Gastropoda, Prosobranchia) in the Pacific Ocean].— Trudy Instituta 
Okeanologii 81:144—-152. 

Barskov, I. S., M. A. Golovinova, and V. N. Goryachev. 1980. [Structure of the nacreous layer of 
deep-water mollusks of the genus Seguenzia (Mollusca, Gastropoda)].—Doklady Akademii 
Nauk SSSR 252(4):1015-1017. [In Russian; translation in Doklady Biological Sciences 252: 
312-314.] 


VOLUME 96, NUMBER 4 751 


Bayer, F. M. 1971. New and unusual mollusks collected by R/V JOHN ELLIOTT PILLSBURY 
and R/V GERDA in the tropical western Atlantic.— Bulletin of Marine Science 21(1):111-—236. 

Belyaev, G. M. 1966. Donnaya fauna naibol’shikh glubin (ul’traabissali) Mirovogo okeana. [The 
benthic fauna of the greatest depths (ultra-abyssal) of the world ocean.].—Izdatel’stvo ‘““Nauka,” 
Moscow, 248 pp. 

Boss, K. J. 1982. Seguenziidae, pp. 973-974.—Jn S. P. Parker (ed.), Synopsis and Classification of 
Living Organisms. Vol. 1. McGraw-Hill, New York. 

Bouchet, P. 1976. Mise en évidence de stades larvaires planctoniques chez des Gastéropodes Proso- 
branches des étages bathyal et abyssal.— Bulletin Muséum National d’Histoire Naturelle, Paris 
(3) 400:947-972. 

Bouchet, P., and A. Warén. 1979. Planktotrophic larval development in deepwater gastropods. — 
Sarsia 64(1—2):37—40. 

Cernohorsky, W. O. 1978. Tropical Pacific marine shells. Pacific Publications, Sidney/New York, 
352 pp. 

Clarke, A. H., Jr. 1959. New abyssal molluscs from off Bermuda collected by the Lamont Geological 

Laboratory. — Proceedings of the Malacological Society of London 33(5):231-238. 

1961. Abyssal mollusks from the South Atlantic Ocean.— Bulletin of the Museum of Com- 
parative Zoology 125(12):345-387. 

. 1962. Annotated list and bibliography of the abyssal marine molluscs of the world. — National 

Museum of Canada, Bulletin 181:114 pp. 

Cossmann, M. 1888. Catalogue illustré des coquilles fossiles de l’Eocéne des environs de Paris. III. — 

Annales de la Société Royale Malacologique de Belgique 23:3-324. 

1918. Essais de Paléoconchologie comparée. Vol. 11. Paris, 388 pp. 

Cotter, G. de P. 1907. Fossils from the Miocene of Burma.—Records of the Geological Survey of 
India, Calcutta 36:131-132. 

Cotton, B. C. 1946. Australian beaded top shells.—South Australian Naturalist 23(3):6—-8. 

1959. South Australian Mollusca. Archaeogastropoda. Handbook of the flora and fauna of 
South Australia. Government Printer, Adelaide, 449 pp. 

, and F. K. Godfrey. 1931. South Australian shells, including descriptions of new genera and 

species.—South Australian Naturalist 12(4):51-63. 

, and 1938. New species of South Australian Gastropoda.—Records of the South 

Australian Museum, Adelaide 6(3):199—206. 

Crisp, M. 1981. Epithelial sensory structures of trochids.—Journal of the Marine Biological Asso- 
ciation of the United Kingdom 61(1):95-106. 

Dall, W. H. 1881. Preliminary report on the Mollusca. Reports on the results of dredging . . . in the 
Gulf of Mexico, and in the Caribbean Sea, 1877-79, by the United States Coast Survey steamer 
“Blake”. ..—Bulletin of the Museum of Comparative Zoology 9(2):33-144. 

—. 1885. List of marine Mollusca comprising the Quaternary fossils and Recent forms from 

American localities between Cape Hatteras and Cape Roque including the Bermudas. — United 

States Geological Survey Bulletin 24:1-336. 

1889a. Notes on the soft parts of Trochus infundibulum Watson, with an account of a 
remarkable sexual modification of the epipodium, hitherto undescribed in Mollusca. — Nautilus 
3:2-4. 

. 1889b. Report on the Mollusca. Part II. Gastropoda and Scaphopoda. Reports on the results 

of dredging .. . in the Gulf of Mexico (1877-78) and in the Caribbean Sea (1879-80), by the 

U.S. Coast Survey steamer “Blake” . . .— Bulletin of the Museum of Comparative Zoology 18: 

1-492. 

—. 1889c. A preliminary catalogue of the shell-bearing marine mollusks and brachiopods of 
the southeastern coast of the United States.— Bulletin of the United States National Museum 

37:1-221. 

1890. Preliminary report on the collection of Mollusca and Brachiopoda obtained in 1887- 
88. Scientific results of explorations by the U.S. Fish Commission steamer “Albatross.” No. 
VII.— Proceedings of the United States National Museum 12:219-362. 

1907. Descriptions of new species of shells, chiefly Buccinidae, from the dredgings of the 
U.S.S. “Albatross’ during 1906, in the northwestern Pacific, Bering, Okhotsk, and Japanese 
Seas. —Smithsonian Miscellaneous Collection 50, pt. 2, no. 1727:139-173. 


V2 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


1908. Reports on the dredging operations off the west coast of Central America to the 
Galapagos... XXXVI. Reports on the scientific results of the expedition to the eastern tropical 
Pacific .. . by the U.S. Fish Commission steamer “Albatross,” from October, 1904, to March, 
1905... XIV. The Mollusca and Brachipoda. — Bulletin of the Museum of Comparative Zoology 
43(6):205—487. 

. 1919. Descriptions of new species of Mollusca from the north Pacific Ocean in the collection 
of the United States National Museum.— Proceedings of the United States National Museum 
56(2295):293-371. 

—. 1921. Summary of the marine shellbearing mollusks of the northwest coast of America, 
from San Diego, California, to the Polar Sea, mostly contained in the collection of the United 
States National Museum, with illustrations of hitherto unfigured species. — Bulletin of the United 
States National Museum 112:1-—217. 

. 1925. Illustrations of unfigured types of shells in the collection of the United States National 
Museum. — Proceedings of the United States National Museum 66(2554):1—41. 

1927. Small shells from dredgings off the southeast coast of the United States by the United 
States Fisheries steamer “Albatross” in 1885 and 1886.—Proceedings of the United States 
National Museum 70(2667):1—134. 

Dautzenberg, P. 1889. Contribution a la faune malacologique des Iles Agores.— Résultats des Cam- 
pagnes Scientifiques accomplies sur son yacht par le Prince Albert I**, Prince de Monaco 1:1— 
112. 

—. 1925. Mollusques nouveaux provenant des croisiéres du Prince Albert I de Monaco.— 
Bulletin de l’Institut Océanographique de Monaco No. 457:1-12. 

—. 1927. Mollusques provenant des campagnes scientifiques du Prince Albert I** de Monaco 

dans |’Ocean Atlantique et dans le Golfe de Gascogne. — Résultats des Campagnes Scientifiques 

accomplies sur son yacht par le Prince Albert I**, Prince de Monaco 72:1—400. 

, and H. Fischer. 1897a. Campagnes scientifiques de S. A. le Prince Albert I** de Monaco. 

Dragages effectués par l’Hirondelle et par la Princesse Alice, 1888—1896.—Memoires de la 

Société Zoologique de France 10:139-234. 


, and 1897b. Campagnes scientifiques de S. A. le Prince Albert I* de Monaco. Di- 
agnoses d’espéces nouvelles de gasteropodes.— Bulletin de la Société Zoologique de France 22: 
37-45. 

, and . 1906. Mollusques provenant des dragages effectués 4 l’ouest de l’Afrique pendant 


les campagnes scientifiques de S. A. S. le Prince de Monaco. — Résultats des Campagnes Scien- 
tifiques accomplies sur son yacht par le Prince Albert I**, Prince de Monaco 32:1—125. 
Erben, H. K. 1972. Uber die Bildung und das Wachstum von Perlmutt.—Biomineralisation 4: 
15-46. 
Fischer, P. 1882. Diagnoses d’espéces nouvelles de mollusques recuillis dans le cours de expéditions 
scientifiques de l’aviso le Travailleur (1880 et 1881).—Journal de Conchyliologie 30:49-55S. 
1885. Manuel de Conchyliologie et de Paléontologie conchyliologique, ou historie naturelle 
de mollusques vivants et fossiles. F. Savy, Paris, pp. 689-896. 
Fretter, V. 1964. Observations on the anatomy of Mikadotrochus amabilis Bayer.— Bulletin of Marine 
Science 14(1):172-184. 
. 1965. Functional studies of the anatomy of some neritid prosobranchs.—Journal of Zoology 


147:46-74. 
. 1966. Observations on the anatomy of Perotrochus.— Bulletin of Marine Science 16(3):603- 
614. 


1975. Umbonium vestiarium, a filter-feeding trochid.—Journal of Zoology 177:541—552. 
, and A. Graham. 1962. British Prosobranch Molluscs. The Ray Society, London, 755 pp. 
, and . 1977. The prosobranch molluscs of Britain and Denmark. Part 2— Trochacea. — 
Journal of Molluscan Studies, Supplement 3:39-100. 
, and 1978a. The prosobranch molluscs of Britain and Denmark. Part 3—Neritacea, 
Viviparacea, Valvatacea, terrestrial and freshwater Littorinacea and Rissoacea.—Journal of 
Molluscan Studies, Supplement 5:101-152. 
, and 1978b. The prosobranch molluscs of Britain and Denmark. Part 4. Marine 
Rissoacea.— Journal of Molluscan Studies, Supplement 6:153-241. 
——., ———, and J. H. McLean. 1981. The anatomy of the Galapagos Rift limpet, Neomphalus 

fretterae.—Malacologia 21(1—2):337-361. 


VOLUME 96, NUMBER 4 753 


Giles, E.,and J. Gosliner. 1983. Primary type specimens of marine Mollusca (excluding Cephalopoda) 
in the South African Museum.— Annals of the South African Museum 92(1):1-—52. 

Golikov, A. N., and Y. I. Starobogatov. 1975. Systematics of prosobranch gastropods. — Malacologia 
15(1):185—232. 

Goryachev, V.N. 1979. K sisteme glubokovodnikh mollyuskov semeistva Seguenziidae (Gastropo- 
da). [On the system of the deep-sea molluscan family Seguenziidae (Gastropoda)]. Pp. 70-71 
in J. M. Likharev (ed.), Molluscs. Main results of their study. Abstracts of communications. 
Zoological Institute, Akademiya Nauk SSSR, Leningrad. 

Graham, A. 1965. Observations on the anatomy of some trochacean gastropods. — Bulletin of Marine 
Science 15(1):202-—210. 

Habe, T. 1953. Descriptions of twelve new Japanese shells. — Venus 17(3):130-144. 

1955a. Notes on the systematic position of the genus Orectospira Dall, 1925.—Zoological 

Magazine 64:259—260. 

—. 1955b. Note on the systematic position of the genus Orectospira Dall, 1925.—Minutes of 
the Conchological Club of Southern California 147:4. 

—. 1961. Coloured illustrations of the shells of Japan. Vol. II. Hoikusha Publ. Co., Osaka, 


183+42 pp. 
1964. Shells of the western Pacific in color. Vol. IJ. Hoikusha Publ. Co., Osaka, [7]+ 
233 pp. 


Harris, G. F. 1897. Catalogue of Tertiary Mollusca in the Department of Geology, British Museum 
(Natural History). I. Australasian Tertiary Mollusca. London, 407 pp. 

Hedley, C. 1905. Mollusca from one hundred and eleven fathoms, east of Cape Byron, New South 

Wales.— Records of the Australian Museum 6(2):41—54. 

1907. The results of deep-sea investigations in the Tasman Sea. 3. Mollusca from eighty 
fathoms off Narrabeen.— Records of the Australian Museum 6(4):283-304. 

, and W. L. May. 1908. Mollusca from one hundred fathoms seven miles east of Cape Pillar, 

Tasmania.— Records of the Australian Museum 7:108-125. 

Hickman, C.S. 1980. Gastropod radulae and the assessment of form in evolutionary paleontology. — 
Paleobiology 6(3):276-294. 

Houbrick, R. S. 1980. Review of the deep-sea genus Argyropeza (Gastropoda: Prosobranchia: Cer- 
ithiidae). —Smithsonian Contributions to Zoology 321:1—30. 

Jeffreys, J. G. 1876. Preliminary report of the biological results of a cruise in H. M. S. ‘Valorous’ 

to Davis Strait in 1875.— Proceedings of the Royal Society of London 25:177-230. 

1877. New and peculiar Mollusca of the Eulimidae and other families of Gastropoda, as 
well as of the Pteropoda, procured in the ‘Valorous’ Expedition.—Annals and Magazine of 
Natural History (4)19:317-339. 

—. 1879. Notes as to the position of the genus Seguenzia among the Gastropoda.—Journal of 
the Linnean Society of London, Zoology 14:605-606. 

——. 1880a. The deep-sea Mollusca of the Bay of Biscay.—Annals and Magazine of Natural 

History (5)6:315-319. 

1880b. The French deep-sea exploration in the Bay of Biscay.—Report of the British As- 
sociation for 1880:1—13. 

1883. Mediterranean Mollusca (No. 3) and other Invertebrata.—Annals and Magazine of 
Natural History (5)11:393—401. 

1885. On the Mollusca procured during the “Lighting? and ‘Porcupine’ Expeditions, 1868- 
70. (Part 9).—Proceedings of the Zoological Society of London for 1885:27-63. 

Johnson, C. W. 1934. List of marine Mollusca of the Atlantic coast from Labrador to Texas.— 
Proceedings of the Boston Society of Natural History 40(1):1—204. 

Jung, P. 1975. Quaternary larval gastropods from Leg 15, Site 147, Deep Sea Drilling Project. 
Preliminary report.— Veliger 18(2):109-126. 

Kanno, S. 1958. New Tertiary molluscs from the Chichibu Basin, Saitama Prefecture, Central 
Japan. —Science Report. Tokyo Bunrika Daig C6, No. 55:157-229. 

Keen, A.M. 1971. Seashells of tropical West America. 2nd Ed. Stanford University Press, Stanford, 

California, 1064 pp. 

, and E. Coan. 1974. Marine molluscan genera of western North America. An illustrated key. 

2nd Ed. Stanford University Press, Stanford, 208 pp. 

, and L. R. Cox. 1960. [Margaritinae]. Pp. 1249-1251 in R. C. Moore (ed.), Treatise on 


754 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Invertebrate Paleontology. Part I, Mollusca 1. Geological Society of America, University of 
Kansas Press. 

Kensley, B. 1973. Sea-shells of southern Africa—Gastropods. Maskew Miller Ltd., Cape Town, 236 
pp. 

Knight, J.B. 1945. Some new genera of Paleozoic Gastropoda. — Journal of Paleontology 19(6):573- 

587. 

, R. L. Batten, and E. L. Yochelson. 1960. [Trochonematacea]. Pp. 1224-1225 in R. C. Moore 

(ed.), Treatise on Invertebrate Paleontology. Part I, Mollusca 1. Geological Society of America, 

University of Kansas Press. 

Knudsen, J. 1964. Scaphopoda and Gastropoda from depths exceeding 6000 meters.—Galathea 
Report 7:125-136. 

Kobelt, W. 1878. Illustrirtes Conchylienbuch. Niirnberg, 331 pp. 

1886-1888. Prodromus Faunae Molluscorum Testaceorum maria Europaea inhabitantium. 
Niirmberg, 550 pp. 

Koperberg, E.G. 1931. Jungtertiare und Quartére Mollusken von Timor. —Jaarboek van het Mijnwezen 
in Nederlandsch-Indié 59:1—-165. 

Kosuge, S. 1966. The family Triphoridae and its systematic position.— Malacologia 4(2):297—324. 

Ladd, H.S. 1970. Eocene mollusks from Eua, Tonga.— United States Geological Survey, Professional 
Paper 640-C:1-111, C1-—C10. 

—. 1982. Cenozoic fossil molluscs from western Pacific islands; Gastropods (Eulimidae and 
Volutidae through Terebridae).— United States Geological Survey, Professional Paper 1171:1- 
100. 

Laubier, L.,and P. Bouchet. 1976. Unnouveaucopépode parasite de la cavité palléale d’un gastéropode 
bathyal dans le Golfe de Gascogne Myzotheridion seguenziae gen. sp. nov.— Archives Zoologie 
Expérimentale et Générale 117(4):469-484. 

Locard, A. 1886. Prodrome de Malacologie francaise. Catalogue général des Mollusques vivants de 

France (Mollusques marins). Lyon, Paris, 779 pp. 

. 1898. Expeditions scientifiques du ““Travailleur” et du “Talisman” pendant les années 1880, 

1881, 1882, 1883. Mollusques testaces. II. Paris, 515 pp. 

1899. Les Coquilles marines au large des Cétes de France. Paris, 198 pp. 

Marshall, B. A. (In press). Recent and Tertiary Seguenziidae (Mollusca: Gastropoda) from the New 
Zealand region.— New Zealand Journal of Zoology. 

Martens, E. von. 1881. Mollusca.—Zoological Record 16, 93 pp. 

. 1901. Neue Meeres-Conchylien von der Deutschen Tiefsee-Expedition. — Sitzungs-Berichten 

der Gesellschaft Naturforschender Freunde Berlin 1901:14-26. 

—. 1903. Die beschalten Gastropoden der Deutschen Tiefsee-Expedition 1898-1899. A. Sys- 
tematisch-geographischer Teil. — Wissenschaftliche Ergebnisse der deutschen Tiefsee Expedition 
auf dem Dampfer “Valdivia.” 7:1-146. 

Maury, C. J. 1922. Recent Mollusca of the Gulf of Mexico and Pleistocene and Pliocene species 
from the Gulf States. Part 2: Scaphopoda, Gastropoda, Amphineura, Cephalopoda. — Bulletins 
of American Paleontology 9(38):34—-142. 

McLean, J. H. 1981. The Galapagos Rift limpet Neomphalus: Relevance to understanding the 
evolution of a major Paleozoic-Mesozoic radiation. — Malacologia 21(1—2):291-336. 

Meek, F. B. 1864. Description of Carboniferous fossils.—Geological Survey of California, Paleon- 
tology of California 1:3-16. 

Melvill, J.C. 1904. Description of twenty-eight species of Gastropoda from the Persian Gulf, Gulf 

of Oman, and Arabian Sea, dredged by Mr. F. W. Townsend, of the Indo-European Telegraph 

Service, 1900—1904.— Proceedings of the Malacological Society of London 6:158-169. 

1910. Descriptions of twenty-nine species of marine Mollusca from the Persian Gulf, Gulf 
of Oman, and North Arabian Sea, mostly collected by Mr. F. W. Townsend, of the Indo- 
European Telegraph Service.— Annals and Magazine of Natural History (8)7:1—17. 

, and R. Standen. 1903. Descriptions of sixty-eight new Gastropoda from the Persian Gulf, 

Gulf of Oman, and North Arabian Sea, dredged by Mr. F. W. Townsend, of the Indo-European 

Telegraph Service, 1901—1903.—Annals and Magazine of Natural History (7)12:289-324. 

Merrill, A. S. 1970a. Fluxina Dall is a Calliostoma Swainson.—Nautilus 84(1):32-34. 

1970b. The family Architectonicidae (Gastropoda: Mollusca) in the western and eastern 
Atlantic.— Unpublished Ph.D. Dissertation, University of Delaware, [6] + 338 pp. 


VOLUME 96, NUMBER 4 755 


Morris, P. A., and W. J. Clench. 1973. A Field Guide to shells of the Atlantic and Gulf Coasts and 
the West Indies. Houghton Mifflin Co., Boston, 330 pp. 

Nobre, A. 1884. Molluscos marinhos de Portugal. Porto. 

1932. Molluscos marinhos de Portugal. Vol. 1. Porto, 466 pp. 

1936. Molluscos marinhos de Portugal. Vol. 2. Porto, 381 pp. 

——. 1938-1940. Fauna malacologica de Portugal: I. Moluscos marinhos e das Aguas salobras. 
Porto, 807 pp. 

Noetling, F. 1901. The fauna of the Miocene beds of Burma.—Palaeontologia Indica, N. S., 1(3), 
378 pp. 

Nordsieck, F. 1968. Die Europaéischen Meeres-Gehauseschnecken (Prosobranchia) vom Eismeer bis 

Kapverden und Mittelmeer. Gustav Fischer, Stuttgart, 273 pp. 

1973. Abyssal mollusks from the Ionic Sea.—La Conchiglia 5(1 1—12):4—-7. 

Odhner, N. H. 1932. Zur Morphologie und Systematik der Fissurelliden.—Jenaische Zeitschrift fiir 
Naturwissenschaft 67:292-309. 

Okutani, T. 1964. Report on the archibenthal and abyssal gastropod Mollusca mainly collected from 
Sagami Bay and adjacent waters by the R. V. ““Soyo-Marw’’ during the years 1955-1963.— 
Journal of the Faculty of Science, University of Tokyo, Sect. II, 15:371-—447. 

—. 1968. Bathyal and abyssal Mollusca trawled from Sagami Bay and the south off Boso 

Peninsula by the R/V “Soyo-Maru,” 1965-1967.—Bulletin of the Tokai Regional Fisheries 

Research Laboratory 56:7—-55. 

. 1974. Review and new records of abyssal and hadal molluscan fauna in Japanese and adjacent 

waters. — Venus 33(1):23-39. 

1982. Rediscoveries of an abyssal trochid, Basilissa superba Watson from the south of 

Japan. — Venus 40(4):237-239. 

Oldroyd, I. S. 1927. The marine shells of the west coast of North America. Vol. 2. Gastropoda, 
Scaphopoda, and Amphineura. Part 2. Stanford University Publications, University Series, 
Geological Sciences, 304 pp. 

Oppenheim, P. 1906. Zur Kenntnis alttertiarer Faunen in Agypten. 2 Lieferung: Der Bivalven zweiter 
Teil, Gastropoda und Cephalopoda.— Paleontographica 30:163-348. 

—. 1923. Uber eine Eocinfauna der Polje von Lukevac bei Nevesinje in der Herzegowina. 
Berlin, 100 pp. 

Pelseneer, P. 1906. Mollusca.—Jn E. R. Lankester (ed.), A Treatise on Zoology. Pt. V. Adam and 
Charles Black, London, 355 pp. 

Perrilliat, M. delC. 1972. Monographia de los moluscos del Mioceno medio de Santa Rosa, Veracruz, 
México. Parte 1. (Gasteropodos: Fissurellidae a Olividae). — Paleontologia Mexicana 32:1-232. 

—. 1974. Catalogo de moluscos del terciario del sur de México (Estados de Veracruz, Oaxaca 
y Chiapas). — Paleontologia Mexicana 38:1-66. 

Philippi, R. A. 1844. Enumeratio Molluscorum Siciliae cum viventium tum tellure tertiaria fossilium, 
quae itinere suo observavit. Vol. 2. Halis Saxonum, 303 pp. 

Popov, S. V., and I. S. Barskov. 1978. Shell structure of mollusks and its value in phylogeny and 
classification. — Malacological Review 11(1—2):152-153. (Abstract of report to 5th Meeting on 
the Investigation of Molluscs, Leningrad, Feb. 1975). 

Porter, H. J. 1974. The North Carolina marine Mollusca—an atlas of occurrence. University of 
North Carolina, Institute of Marine Sciences, Morehead City, North Carolina, 351 pp. 
Pulley, T. E. 1952. An illustrated check list of the marine mollusks of Texas.—Texas Journal of 

Science 2:167-199. 

Quinn, J. F., Jr. 1979. Biological results of the University of Miami Deep-Sea Expeditions. 130. 

The systematics and zoogeography of the gastropod family Trochidae collected in the Straits 

of Florida and its approaches. — Malacologia 19(1):1-62. 

1981. A preliminary overview of the Seguenziidae Verrill, 1884.— Bulletin of the American 

Malacological Union for 1980:74 [abstract]. 

1983a. The Seguenziacea: An update.—American Malacological Bulletin 1(1). [Abstract]. 
1983b. Carenzia, a new genus of Seguenziacea (Gastropoda: Prosobranchia) with the de- 
scription of a new species from the northeastern Pacific. — Proceedings of the Biological Society 

of Washington 96:355-364. 

. In preparation. A revision of the Seguenziacea Verrill, 1884 (Gastropoda: Prosobranchia). 

II. Systematics of the western Atlantic species. 


756 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Ravn, J. P. J. 1933. Etudes sur les Pélécypodes et gastropodes daniens du Calcaire de Faxe.— 
Kongelige Danske Videnskabernes Selskabs Skrifter (9)5(2):3-74. 
—. 1939. Etudes sur les Mollusques du Paléocéne de Copenhague.—Biologiske Skrifter utgitt 
av Det Kongelige Danske Videnskabernes Selskabs 1(1):1—106. 
Rex, M. A. 1972. Species diversity and character variation in some western North Atlantic deep 
sea gastropods. — Unpublished Ph.D. Dissertation. Harvard University, 178 pp. 
—. 1976. Biological accommodation in the deep-sea benthos: comparative evidence on the 
importance of predation and productivity.— Deep-Sea Research 23(10):975-987. 
Riedel, L. 1932. Die Oberkreide vom Mungofluss in Kamerun und ihre Fauna.—Beitrage zur geo- 
logischen Erforschung der deutschen Schutzgebeite, 16:1—154. 
Rios, E. C. 1975. Brazilian marine mollusks iconography. Fundacao Universidade do Rio Grande, 
Rio Grande-RS, 331 pp. 
Rokop, F. J. 1972. Notes on abyssal gastropods of the eastern Pacific, with descriptions of three new 
species. — Veliger 15(1):15-19. 
Schepman, M. M. 1908. The Prosobranchia of the Siboga Expedition. Part 1. Rhipidoglossa and 
Docoglossa. —Siboga Expedition, Monographie 49!a:1-107. 
. 1909. The Prosobranchia of the Siboga Expedition. Part 2. Taenioglossa and Ptenoglossa. — 
Siboga Expedition, Monographie 49'b:109-231. 
Seguenza, G. 1876. Studii stratigrafici sulla Formazione pliocenica dell’Italia Meridionale. Elenco 
dei Cirripedi e dei Molluschi della zona superiore dell’antico plioceno.— Reale Comitato Geo- 
logico Italiana, Bolletino 7(5—6):180-189. 
1877. Brevissimi cenni intorno le Formazioni Terziarie della provincia di Reggio-Calabria. 
Messina, 31 pp. 
1879. Le formazione Terziarie nella provincia di Reggio (Calabria). — Atti Reale Accademia 
dei Lincei, Memorie (3)1:1—446. 
Shimek, R. L., and A. J. Kohn. 1981. Functional morphology and evolution of the toxoglossan 
radula.— Malacologia 20(2):423—438. 
Sorgenfrei, F. 1958. Molluscan assemblages from the marine Middle Miocene of South Jutland and 
their environments.— Danmarks Geologiske Undersogelse (II) 79:1-503. 
Tate, R. 1888. Census of the fauna of the older Tertiary of Australia.— Journal and Proceedings of 
the Royal Society of New South Wales 22:240-253. 
1890. The gastropods of the older Tertiary of Australia.— Transactions of the Royal Society 
of South Australia 13:185-235. 
—. 1894. Unrecorded genera of the older Tertiary fauna of Australia.—Journal of the Royal 
Society of New South Wales 27:169-191. 
Taylor, D. W., and N. F. Sohl. 1962. An outline of gastropod classification.—Malacologia 1(1): 
7-32. 
Thiele, J. 1903. Die beschalten Gastropoden der Deutschen Teifsee-Expedition 1898-1899. B. An- 
atomisch—systematische Untersuchungen einiger Gastropoden. — Wissenschaftliche Ergebnisse 
der deutschen Tiefsee Expedition auf dem Dampfer “‘Valdivia” 7:147-180. 
1925. Gastropoda der Deutschen Tiefsee-Expedition. II. — Wissenschaftliche Ergebnisse der 
deutschen Tiefsee Expedition auf dem Dampfer “Valdivia” 17(2):36-382. 
——. 1929-35. Handbuch der systematischen Weichtierkunde. 2 vols. Gustav Fischer, Jena, 
1134 pp. 
Treece, G. D. 1977. Bathymetric records of marine shelled Mollusca from the northern shelf of 
Yucatan, Mexico.— Unpublished M.S. Thesis, Texas A&M University, 212 pp. 
—. 1979. Living marine mollusks from the south Texas continental shelf.—Texas Journal of 
Science 31(3):271-283. 
. 1980. Bathymetric records of marine shelled Mollusca from the northeastern shelf and upper 
slope of Yucatan, Mexico.— Bulletin of Marine Science 30(3):552—570. 
Tryon, G. W. 1883. Structural and systematic conchology: An introduction to the study of the 
Mollusca. Vol. 2. Philadelphia, 430 pp. 
—. 1887. Manual of Conchology. Vol. 9. Solariidae, Ianthinidae, Trichotropidae, Scalariidae, 
Cerithiidae, Rissoidae, Littorinidae. Philadelphia, 488 pp. 
Verco, J.C. 1906. Notes on South Australian marine Mollusca, with descriptions of new species— 
Part IV.—Transactions and Proceedings and Report of the Royal Society of South Australia 
30:205-224. 


VOLUME 96, NUMBER 4 757 


Vermill, A. E. 1884. Second catalogue of Mollusca recently added to the fauna of the New England 

coast and the adjacent parts of the Atlantic, consisting mostly of deep-sea species, with notes 

on others previously recorded.— Transactions of the Connecticut Academy of Science 6:139- 

294. 

1886. Results of the explorations made by the steamer “‘Albatross,”’ off the northern coast 
of the United States in 1883.—Report of the Commissioner of Fish and Fisheries for 1883: 
503-699. 

Warén, A. 1980. Marine Mollusca described by John Gwyn Jeffreys, with the location of the type 
material.—Conchological Society of Great Britain and Ireland, Special Publication 1:1-60. 

Watson, R. B. 1879a. Mollusca of H.M.S. ‘Challenger’ Expedition. III. Trochidae, viz. the genera 

Seguenzia, Basilissa, Gaza and Bembix.—Journal of the Linnean Society of London, Zoology 

14:586-605. 

. 1879b. Mollusca of H.M.S. ‘Challenger’ Expedition. IV. Trochidae continued, viz. the genera 

Basilissa and Trochus, and the Turbinidae, viz. the genus Turbo.—Journal of the Linnean 

Society of London, Zoology 14:692-716. 

1886. Report on the Scaphopoda and Gasteropoda collected by H.M.S. Challenger during 
the Years 1873-76.—Report on the Scientific Results of the Voyage of H.M.S. Challenger, 
1873-1876, Zoology 15:1-680. 

Wenz, W. 1938. Gastropoda. Allgemeine Teil und Prosobranchia.— Jn O. Schindewolf, Handbuch 
der Paléozoologie, Band 6, Teil 1, Leif. 1 & 2:1—480. 

Wise, S. W., Jr. 1970. Microarchitecture and mode of formation of nacre (mother-of-pearl) in 
pelecypods, gastropods and cephalopods.—Eclogae Geologicae Helvetiae 63:775-797. 
Woodring, W.P. 1928. Miocene mollusks from Bowden, Jamaica. Part II. Gastropods and discussion 

of results.— Carnegie Institution of Washington, Publication 385:1-564. 

Yochelson, E. L. 1956. Permian Gastropoda of the southwestern United States: I. Euomphalacea, 
Trochonematacea, Pseudophoracea, Anomphalacea, Craspedostomatacea, Platyceratacea. — 
Bulletin of the American Museum of Natural History 110(3):173-276. 

Yokoyama, M. 1922. Fossils from the Upper Murashino of Kazusa and Shimosa.—Journal of the 
College of Science, Tokyo 44:1—200. 


Florida Department of Natural Resources, Bureau of Marine Research, 100 
Eighth Avenue S.E., St. Petersburg, Florida 33701. 


NOTE: While this paper was in press, the monograph cited as “Marshall in press’”’ was published 
(New Zealand Journal of Zoology 10:235—262, September 1983). In that report, Marshall recognizes 
22 new species in eight genera, five of which were new. Since Marshall provided names for Seguenzia 
Groups II and IV (Seguenziella and Fluxinella, respectively), and modified the composition of several 
other taxa of my Table I, the reader is urged to consult his paper for further information. 


PROC. BIOL. SOC. WASH. 
96(4), 1983, pp. 758-769 


OBSERVATIONS ON SPECIES OF THE FOSSIL 
GENUS AXOPORA (COELENTERATA: HYDROZOA) 
AND ITS EVOLUTIONARY SIGNIFICANCE TO THE 

STYLASTERIDAE 


Stephen D. Cairns 


Abstract.—Two species of Axopora are redescribed and the remaining valid 
species are diagnosed and discussed. Three species are illustrated by scanning 
electron photomicrographs. Special emphasis is given to the morphology of the 
gastrostyles, and their supposed homology to those of the stylasterids is discussed. 
A hypothetical evolutionary scheme is proposed suggesting the evolution of the 
axoporids from a hydractiniid ancestor by the acquisition of calcification and the 
transformation of its protective spines to gastrostyles. An ancestor to the axoporids 
is suggested to have evolved into the stylasterids also, the major changes being 
the encasement of its gonozooids and dactylozooids in specialized calcareous 
structures. The Axoporidae is considered to be a family of athecate hydroids with 
close affinities to the Hydractiniidae and Stylasteridae. 


The Axoporidae is a small family of calcified hydrozoans (13-14 nominal species 
in 1-2 genera), known only from the Oligocene to Eocene of Europe and South 
West Africa. They were originally described as Scleractinia (Pocillopora of De- 
france (1826), and Holaraea of Milne Edwards and Haime (1849)) and some were 
even classified as tabulate corals and sponges (A/veolites and Geodia of Michelin 
(1844)). Michelin (1844) also identified a species as a milleporid, and, until 1963, 
most axoporids were assigned to the order Milleporina. Boschma (1963b) removed 
the axoporids from the Milleporina and created a new order, the Axoporina, for 
them, which, according to him, had closer affinities to the Stylasteridae than to 
the Milleporina. 

The classification and phylogenetic position of the axoporids are based strongly 
on the characteristics of their gastrostyles. Until 1963, all axoporid gastrostyles 
were considered to be longitudinally grooved, nonspinose, fasciculate structures. 
However, Boschma (1963b) revealed that they are deeply ridged spinose cylinders, 
not at all fasciculate. Scanning electron microscopy allows an even more detailed 
analysis of gastrostyle morphology and a detailed comparison to other stylasterid 
gastrostyles. 

Following a redescription of two species of Axopora and diagnoses of the other 
valid species, the interrelationships of the axoporids, stylasterids, and athecate 
hydroids will be discussed. 


Class Hydrozoa 


Order Hydroida 
Family Axoporidae Boschma, 1951 


Diagnosis.—Calcified hydrozoans with long spinose gastrostyles and with no 
skeletal evidence of dactylozooids (dactylopores) or gonophores (ampullae). Eocene 


VOLUME 96, NUMBER 4 759 


to Oligocene (?Miocene): Europe, South West Africa, (7New Zealand). One or two 
genera. 


Genus Axopora Milne Edwards and Haime, 1850 


Diagnosis.—Characteristics of the family. Type-species: Geodia pyriformis 
Michelin, 1844. Five to seven species. 


Axopora solanderi (Defrance, 1826) 
Figs. 1-7 


Pocillopora solanderi Defrance, 1826:48. 

Palmipora solanderi.— Michelin 1844:166, pl. 45, fig. 9. 

Lobopora solanderi.— Milne Edwards and Haime 1850:lix. 

Axopora solanderi.— Milne Edwards and Haime 1851:151; 1857:243-244, pl. F3, 
fig. 2.—Boschma 1951:25, fig. 3; 1961:F94, fig. 78-1, 2; 1963a: 107, figs. 2a— 
b, d; 1963b:122, 125, pl. 1, fig. 1, pl. 2, figs. 1-2, pls. 3-8. 

Axopora michelini Duncan, 1866:50, pl. 7, figs. 11-15.—Boschma 1963b:124, 
126. 

Axopora fisheri Duncan, 1866:64, pl. 10, figs. 20—22.— Boschma 1963b:124-125, 
26. 

Millepora mamillosa d’Archiardi, 1867:11. 

Axopora mamillosa.—Steinmann 1903:5.—Boschma 1963b:125-126. 


Description.—Corallum composed of flattened lobes (typical form) (Figs. 1—2) 
or lumpy masses (michelini form), but never forming laminar encrustations. In 
both lumpy and lobate forms, gastrostyles originate on both sides of a midline 
(Fig. 3), a plane in center of lobe or lump, not differing in porosity from rest of 
corallum. Surface coenosteum usually smooth but may be ridged, especially at 
tips of lobate specimens. Coenosteal ridges subparallel and roughly perpendicular 
to lobe edge, up to | mm tall and 1.5 mm broad. Lower relief ridges sometimes 
cross between adjacent parallel ridges producing reticulate pattern (Fig. 1). Coe- 
nosteal texture appears to be reticulate-granular, but preservation of specimens 
examined did not allow a definitive categorization. 

Gastropores round, 0.15—0.35 mm in diameter, flush with coenosteal surface 
(Fig. 5). Gastropores randomly arranged, occurring at a density of 1.6—-2.2 pores 
per mm7?. Very diffuse ring palisade occurs in upper part of gastropore tube, ele- 
ments measuring about 25 wm in diameter. Gastrostyles extend almost to coe- 
nosteal surface and occupy most of space within gastropores, unlike in the sty- 
lasterids, where there is always considerable space around the style. 

Length of gastrostyles and number of tabulae per gastrostyle vary with width 
of lobe or lump in which they occur (Figs. 4, 7). In thick coralla, gastrostyles up 
to 3.8 mm long and have over 18 tabulae. Styles rarely exceed 0.21 mm in 
diameter; maximum H:W observed was 23.5. According to Boschma (1963b: 
121), gastrostyles sharply ridged, with spines occurring on ridges; however, scan- 
ning electron microscopy reveals that base of styles definitely not ridged, but 
spinose. Above base of style, spines vertically aligned, sometimes 1-3 across, and 
separated from adjacent rows by shallow groove. Spines so long, and sometimes 
bifurcate, that it is difficult to distinguish actual ridges on style (Figs. 6, 7). Gas- 
trostyle spines cylindrical, up to 0.13 mm long and about 30 um in diameter. 


760 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Figs. 1-6. Axopora solanderi from Valmondois (USNM 68437). 1-2, distal lobes, x1.8, 1.3, 
respectively; 3, Cross section of lobe showing midline and gastrostyles, x 6; 4, A gastrostyle and tabulae 
from colony of Fig. 3, x27; 5, Coenosteum and gastropores, x40; 6, Detail of gastrostyle, x 250. 


VOLUME 96, NUMBER 4 761 


EIS 


> 


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iF tee eS 


es 


2S as, 


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weet 


Figs. 7-9. Gastrostyles. 7, Stereo view of gastrostyle and tabulae of Axopora solanderi, British 
Museum R 50812, x 140; 8, Gastrostyle and tabulae of Sporadopora dichotoma from the Drake Passage 
(Hero 715-895), x 100; 9, Gastrostyle tip of Lepidopora granulosa from Drake Passage (Eltanin sta 


740), x 190. 


Tabulae invariably present, occurring at irregular intervals, averaging about every 
0.25 mm. Tabulae about 6 um thick, completely sealing off lower gastropore tube 
from upper tube. 

Discussion.—Axopora solanderi is distinguished from the other species in the 
genus by its nonencrusting growth form and massive corallum. Its gastrostyles 
are superficially similar to those of Sporadopora dichotoma, particularly with 
regard to the high H:W, similar tabulae, and apparent ridges (Fig. 8); however, 
the gastrostyles of S. dichotoma are distinctly ridged and bear much smaller and 
sharper spines. The characteristics of the tabulae are quite variable and not usually 
considered to be of generic or even specific value. Of all the stylasterines, Lepi- 
dopora granulosa (Fig. 9) has gastrostyles most similar to those of A. solanderi. 

Distribution. —Eocene to Oligocene: England, France, Italy. 


762 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Material examined.— Auvers-sur-Oise, Seine et Oise, France, 7 fragments 
(michelini form), USNM 80889 (Invertebrate Paleontology Series); Valmondois, 
France, 2 fragments (typical form), USNM 68437; Sands of Beauchamps, Auvers, 
France, 2 fragments (michelini form), USNM 68438; Parnes, Beavoir, France, 1 
fragment (michelini form), USNM 80891 (Invertebrate Paleontology Series); Le 
Bois Gouet, Lutecien, France (typical form), USNM 68439; Paris Basin, France, 
1 fragment (michelini form), British Museum R 50812. 


Axopora parisiensis (Michelin, 1844) 
Figs. 10-15 


Alveolites parisiensis Michelin, 1844:166, pl. 45, fig. 10. 

Holaraea parisiensis.— Milne Edwards and Haime 1849:259; 1850:1vi, 40, pl. 7, 
fig. 2. 

Axopora parisiensis.— Milne Edwards and Haime 1851:151; 1857:244.—Bosch- 
ma 1951:25; 1963a:107-109, fig. 1; 1963b:122-123, 126. 


Description. —Corallum always encrusts an elongate cylindrical object, such as 
a gorgonian stem, which, when the stem is lost, produces a cylindrical corallum 
with a hollow center (Figs. 10-11). Largest specimen examined 14 mm long and 
6.5 mm in diameter, with hollow center 2.5 x 2.0 mm in diameter. Coenosteal 
encrustation always thin, with no side branches or lobes. Coenosteum papillose 
(Fig. 13), producing spongy coenosteal texture. 

Gastropores round, 0.45—0.55 mm in diameter, surrounded by round to po- 
lygonal, infundibuliform calices (Fig. 12). Otherwise constant-diameter cylindrical 
gastropore becoming flared near coenosteal surface. Calices closely packed in 
honeycomb arrangement, each calice measuring about 1.2 mm in diameter. The 
density of gastropores therefore less than one pore per mm2?. Diffuse ring palisade 
present, similar to that of A. solanderi. Tips of gastrostyles extending only to base 
of funnel-shaped section of gastropore and thus each occupying only about one- 
half of gastropore (Figs. 14-15). 

Because the corallum is never thick, gastrostyles are relatively short and rarely 
have tabulae. A large gastrostyle is 0.9 mm long and 0.18 mm in diameter 
(H:W = 5); however, H:W ratios usually lower than 5. Spination of gastrostyle is 
similar to that of A. solanderi, with cylindrical spines up to 1.25 mm long and 
19-26 wm in diameter arranged in poorly defined vertical rows. The “ridges” of 
these styles are virtually nonextant. 

Discussion. —Axopora parisiensis is easily distinguished by its large infundibuli- 
form calices and its cylindrical encrusting growth form. 

Distribution.— Eocene: France and England. 

Material examined.—Vandane, France, 3 fragments, USNM 68440; Parnes, 
France, 2 fragments, USNM 68441; Campbon, France, 15 fragments, USNM 
68442; Cuise, France, 2 fragments, USNM 68443; St. Lucien de la Haye, France, 
1 fragment, USNM 68444; Grignon, Seine-et-Oise, France, 35 fragments, USNM 
68445. 


Axopora kolosvaryi (Boschma, 1954) 
Figs. 16-17 


Axopora ramea.—Kolosvary 1949:189, pl. 20, fig. 2. 
Axoporella kolosvadryi Boschma, 1954:101-103, figs. 2a—c; 1956:F-94, fig. 78-3a— 
c; 1963a:114-115. 


VOLUME 96, NUMBER 4 763 


Figs. 10-15. Axopora parisiensis from Parnes (USNM 68441). 10, encrusting piece of colony, x 9; 
11, Cross section of encrustation, x13; 12, Infundibuliform calice, x50; 13, Papillose coenosteal 
texture within a calice, x 385; 14-15, Gastrostyle, x 230, x60, respectively. 


764 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Axopora kolosvadryi.—Boschma 1963b:127. 


Diagnosis.—Corallum encrusting (Boschma 1963b:127). Gastropores round, 
about 0.35 mm in diameter, flush with surface. Gastrostyle 80—90 um in diameter, 
extending almost to surface of coenosteum. Style covered with short clawlike 
spines, curved both upwards and downwards on style (Fig. 16). Spines only about 
35 um long, tapering to point from broad base up to 20 wm in diameter (Fig. 17). 
Spines not aligned in rows; gastrostyles not ridged. Tabulae present. 

Discussion. —Axopora kolosvaryi is distinguished from the other species of Ax- 
opora by its very short clawlike gastrostyle spines, which are not arranged in rows. 
Only one tiny specimen was available for examination, inadequate for a rede- 
scription of the species. 

Distribution. —Lower Eocene: Hungary. 

Material examined.—One fragment from Gant, Hungary (topotypic), USNM 
68447. 


Axopora pyriformis (Michelin, 1844) 


Geodia pyriformis Michelin, 1844:178, pl. 46, fig. 2. 

? Millepora parasitica Catullo, 1856:79, pl. 18, fig. 4. 

Axopora pyriformis. — Milne Edwards and Haime 1850:1ix; 1857:244.— Boschma 
1951:25; 1956:F94; 1963a:123, 126, pl. 1, figs. 2-5, pl. 2, figs. 3-4. 


Diagnosis.—Corallum always encrusting, usually on gastropod shells. Gastro- 
pores 0.25—0.40 mm in diameter, occurring with a density of 1.4-1.5 per mm”. 
Otherwise similar to 4. solanderi. 

Discussion. —Boschma (1963b:126) noted that the only difference between A. 
pyriformis and A. solanderi was in growth form, which may in itself be a variable 
character. This allows for the possibility that A. pyriformis is a junior synonym 
of A. solanderi. Unfortunately, no specimens of A. pyriformis were available for 
study. 

Distribution. —Eocene: France. 


Axopora arborea Keferstein, 1859 


Axopora arborea Keferstein, 1859:381, pl. 15, fig. 9.—Boschma 1951:26; 1963b: 
124, 126: 

? Axopora paucipora Keferstein, 1859:382, pl. 15, fig. 10.—Boschma 1963b:124, 
2S. 

Axopora ramea @’Archiardi, 1867:11.—Not Kolosvary 1949:189 (=A. kolosvar- 
yi).—Boschma 1963b:125, 126. 


Diagnosis.—Corallum delicately branched; branches 3—6 mm in diameter. De- 
tails of gastrostyles unknown. 

Discussion.— The poorly known 4. arborea is distinguished from other species 
by its delicately branching growth form. No specimens were available for study. 

Distribution. —Oligocene: Germany. 


Axopora Cleithridium (Squires, 1958), new combination 


Sporadopora cleithridium Squires, 1958:25-27, pl. 1, figs. 8-12. 
Axoporella cleithridium.—Squires, 1962:136—137. 


VOLUME 96, NUMBER 4 765 


Figs. 16-17. Axopora kolosvaryi from Gant, Hungary (USNM 68447), details of gastrostyle spi- 
nation, x310, x925, respectively. 


Diagnosis.—Corallum with thick branches, up to 14 mm in diameter. Gastro- 
pores irregular in shape, 0.1—-0.2 mm in diameter. Gastrostyles up to 5 mm long; 
tabulae present. 

Discussion.—This is the only known Axopora with thick branches. Squires 
(1958) originally described it as a stylasterine, Sporadopora, and even described 
its dactylopores. When he (Squires 1962) later transferred it to Axoporella he did 
not mention the lack of dactylopores. Those referred to in 1958 were probably 
the superficial expression of coenosteal canals. 

Only one specimen was available for study, a paratype from NZGS 5170. It 
does not appear to have dactylopores nor does it have any gastrostyles or tabulae. 
Perhaps the lack of styles and tabulae in this particular specimen is a result of 
poor preservation, but it does cast doubt on the placement of this species in the 
Axoporidae. Its occurrence in the Kapitean (Upper Miocene) of New Zealand 
would make it by far the youngest member of the family. 

Apparently Boschma (1963b) overlooked this species in his revision of Axopora. 

Distribution.— Upper Miocene: New Zealand. 

Material examined. —Whata, Rowallon, S. D., Southland, New Zealand, NZGS 
5170, USNM 68446 (Paratype). 


Other Species 


Axopora microspora and 4A. alpina, both originally described by d’Orbigny 
(1850) in the genus Holaraea, were considered as species dubiae by Boschma 
(1963b) because of insufficient data. Both species are known from Faudon, France. 

Diamantopora lotzi Weissermel, 1913, is the only species in its genus, and 
Diamantopora is the only other genus in the Axoporidae. It was distinguished 
from Axopora by its nongrooved gastrostyles (Boschma 1956); however, this 
characteristic was subsequently shown to be variable within Axopora. Diaman- 
topora lotzi may therefore represent a seventh valid species of Axopora, but until 
specimens can be examined in detail, its position will remain in doubt. It is known 
only from the Eocene of South West Africa. 


766 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Discussion 


Although traditionally allied with the milleporids (from 1844 to 1963) several 
authors had noted the similarities of axoporids and stylasterids, particularly with 
regard to their gastrostyles. RGmer (1863) was perhaps the first to point this out 
when he observed the similarity of the gastrostyle of the fossil stylasterine Den- 
dracis multipora (subsequently synonymized with A/lopora compressa) with those 
of Axopora arborea. Reuss (1865) made a similar observation based on the same 
species. Steinmann (1903) was the foremost advocate of this point of view when 
he remarked that Axopora was undoubtedly the ancestor of the three Recent 
stylasterid genera Sporadopora, Errina (now Errina, Lepidopora, Lepidotheca, 
and Inferiolabiata) and Spinopora (now Stellapora). Boschma (1954) mentioned 
the resemblance of the gastrostyles of A. kolosvaryi with those of other stylasterids 
and finally he (Boschma 1963b) removed the axoporids from the Milleporina, 
creating a separate order for them, the Axoporina. He stated that the axoporids 
were more closely related to the stylasterids than to the milleporids because of 
their similar gastrostyles, and noted that the gastrostyles of Distichopora were 
most similar to those of Axopora. 

I agree that the axoporids are closely related to the stylasterids, but most closely 
to the genus Lepidopora. Points of similarity and presumed homology between 
Lepidopora and Axopora include: 1) a structurally similar gastrostyle; including 
size, H:W, and spination, 2) a diffuse ring palisade, 3) a calcium carbonate skeleton 
with anastomosing coenosteal canals, and 4) a random (nonlinear) arrangement 
of gastropores on the corallum. Points of difference are that Axopora: 1) lacks 
dactylopores, 2) lacks ampullae, 3) has a less spacious gastropore, 4) has tabulae, 
5) has a different coenosteal texture, and 6) has both an encrusting and branching 
growth form. The major differences are the first two: that Axopora lacks dacty- 
lopores and ampullae; however, this does not necessarily mean that Axopora 
lacked dactylozooids or gonozooids, only that it lacked skeletal evidence of these 
features. For instance, the Recent calcified hydroid Janaria has both dactylozooids 
and gonozooids but has no skeletal evidence of these structures because they are 
composed of tissue that projects above the coenosteum, as in Hydractinia and 
most other athecate hydroids. Thus, Axopora may well have had dactylozooids 
and gonozooids but they were not housed in specialized calcareous structures. 
Most assuredly Axopora did have gonozooids, which were probably superficial 
structures as in Hydractinia. 

In a phylogenetic analysis of the stylasterine genera (Cairns, in press), I chose 
the athecate hydroid Hydractinia as the out-group, or sister taxon, to the stylas- 
terine corals. The most plesiomorphic (primitive) genus of stylasterine resulting 
from this analysis was Lepidopora, particularly L. granulosa. Axopora, because 
of its lack of skeletal evidence of dactylozooids and gonozooids and its encrusting 
growth form for some species, shows similarities to Hydractinia. On the other 
hand, of all the stylasterines, Axopora is most similar to Lepidopora, the most 
primitive stylasterine genus. I therefore propose that Axopora evolved from a 
hydractiniid ancestor in the early Tertiary (?Oligocene) and that a common ances- 
tor gave rise to the stylasterines through Lepidopora. Furthermore, I consider the 
axoporids to be a family of athecate hydroids closely related to the Stylasteridae 
and Hydractiniidae. 


VOLUME 96, NUMBER 4 767 


Lo 
eS & § 
oO 
€ o e é 
x WS 
Oo a 
BN xe © 
~N 


x reese 


shorter dactylopores 


branching coralla 


increasing coarelipniey of 
gastro- and dactylopores 


branching coralla 


gastropore capacious gastropore tube 


tubulae 


distinctive coenosteal textures: 
inear-imbricate, reticulate—granular 


distinctive 
coenostea 
texture 


ampullae 


dactylopores (long) 


B 


spines become gastrostyles 
gastropore with ring palisade 


calcified skeleton 


Fig. 18. Suggested evolution of the axoporids and stylasterids indicating significant character state 
changes. A and B are hypothetical ancestors. 


As a hypothetical scenario I suggest the following (Fig. 18). In the Paleocene 
or Late Cretaceous a hydractiniid ancestor (A), which was encrusting and already 
had the gastrostyle homolog in the form of a chitinous spine, evolved the ability 
to calcify its skeleton and to protect its gastrozooids by enclosing them in rudi- 
mentary tubes: the gastropores. The chitinous spine transformed into the calcar- 
eous gastrostyle. The resultant hypothetical ancestor (B), with little further mod- 
ification became the stem that resulted in Axopora. This evolutionary line developed 
a distinctive coenosteal texture, tabulae to stabilize its long gastrostyles, and, for 
some species, the ability to take advantage of the branching mode. Axopora 
became extinct in the Eocene (? Miocene), perhaps because it did not develop the 
further safeguards characteristic of the stylasterines: a protected gonozooid (am- 
pullae) and a protected dactylozooid (dactylopores). (Stylasterines eventually 
evolved quite elaborate structures to protect their zooids, including fixed and 
hinged gastropore lids and dactylopore spines). In addition to the dactylopores 
and ampullae, the line leading to the stylasterines also evolved distinctive coe- 
nosteal textures (imbricate platelets and reticulate-granular coenosteum, among 
others), a more capacious gastropore tube, and an exclusively branching form. 
The characteristic of gastropore tabulae was probably independently acquired by 
several genera, such as Sporadopora, Distichopora, and Errina, and is therefore 
considered a convergent character. 

The fossil record produces an apparent contradiction to the proposed scenario 
in that the earliest known stylasterids occur before the earliest known axoporids: 
the Paleocene (Danian of Denmark) and Eocene, respectively. As a test of my 
hypothesis, but not necessarily a confirmation, I predict the eventual discovery 
of Axopora from the Paleocene. An alternative explanation would be that the 


768 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


axoporids evolved through an intermediate Paleocene ancestor, which did not 
evolve Axopora until the Eocene and for which there is no known fossil record. 


Literature Cited 


d’Archiardi, A. 1867. Coralli fossili del terreno nummulitico dell’Alpi Venete. Catalogo delle specie 
e breve note. Pisa. 
Boschma, H. 1951. Notes on Hydrocorallia.—Zoologische Verhandelingen, Leiden 13:49 pp., 2 pls. 
1954. De famile Axoporidae.—Verslag van de Koninklijke Nederlandse Akademie van 
Wetenschappen 63(4):99-104, 2 text-figs. 
1956. Milleporina and Stylasterina. Pp. F90—F106, text-figs. 75-85 in R. C. Moore (ed.), 
Treatise on Invertebrate Paleontology. Part F. Coelenterata.— University of Kansas Press, Law- 
rence, Kansas. 
—. 1963a. The generic name Axopora.— Proceedings Koninklijke Nederlandse Akademie van 
Wetenschappen (B)66(3):107—117, 2 figs. 
—. 1963b. Notes on species of the genus Axopora.—Proceedings Koninklijke Nederlandse 
Akademie van Wetenschappen (B)66(3):118-129, 8 pls. 
Cairns, S. D. [in press.] A generic revision of the Stylasteridae. Part 2. Phylogenetic revision. — 
Bulletin of Marine Science 34(2). 
Catullo, T. A. 1856. Dei terreni di sedimento superiore delle Venezie e dei fossili Bryozoari, An- 
thozoari e Spongiari ai quali danno ricetto. Padova, 88 pp., 19 pls. 
Defrance, M. J. L. 1826. Pocillopore (Foss.). P. 48 in F. G. Lavrault (ed.), Dictionnaire des Sciences 
Naturelles 42. Strasbourg and Panis. 
Duncan, P. M. 1866. A monograph of the British fossil corals. Part 1. Introduction; Corals from 
the Tertiary formations. — Palaeontographical Society, London, 111 + 66 pp., pls. 1-10. 
Keferstein, W. 1859. Die Korallen der norddeutschen Tertiargebilde.— Zeitschrift der Deutschen 
geologischen Gesellschaft 11:354—-383, pls. 14-15. 

Kolosvary, G. 1949. The Eocene corals of the Hungarian Transdanubian Province.—Fdéldtani K6- 
zlény, Bulletin de la Société Geologique de Hongrie 78:141-—242, pls. 1-22. 

Michelin, J. L.H. 1844. Pp. 149-178, pls. 43-46 in Iconographie Zoophytologique. C. P. Bertrand, 
Paris. 

Milne Edwards, H., and J. Haime. 1849. Mémoire sur les Polypiers appartenant aux groupes naturels 
des Zoanthaires perforés des Zoanthaires tabulés.—Compte Rendu hebdomadaire des Séances 
de l’ Académie des Sciences, Paris 29:257-263. 


, and 1850. A monograph of the British fossil corals. Part 1. Introduction. Corals 
from the Tertiary and Cretaceous formations. —Palaeontographical Society, London, Ixxxv + 
71 pp., 11 pls. 

, and 1851. Monographie des Polypiers fossiles des terraines palaeozoiques, précédée 


d'un tableau général de la classification des Polypes.— Archives du Muséum National d’ Histoire 

Naturelle, Paris 5:502 pp., 20 pls. 

, and 1857-1860. Histoire Naturelle des Coralliaires ou Polypes proprement dits. 

Volume 2, 633 pp. (1857), atlas, 31 pls. (1860). Roret, Paris. 

d@Orbigny, A. D. 1850. Prodrome de Paléontologie stratigraphique universelle des Animaux Moll- 
usques et Rayonnés 2. V. Masson, Paris. 

Reuss, A. E. 1865. Zur Fauna des deutschen Oberoligocans. II. Anthozoen.—Sitzungsberichte der 
Akademie der Wissenschaften in Wien 50:614—690, pls. 6-15. 

Romer, F. A. 1863. Beschreibung der norddeutschen tertiéren Polyparien.— Palaeontographica 9: 
199-245, pls. 35-39. 

Squires, D. F. 1958. The Cretaceous and Tertiary corals of New Zealand.—New Zealand Geological 

Survey Paleontology Bulletin 29:105 pp., 16 pls. 

1962. Additional Cretaceous and Tertiary coral from New Zealand.—Transactions of the 

Royal Society of New Zealand 1(9):133-150, 4 pls. 

Steinmann, G. 1903. Nachtraége zur Fauna von Stramberg. II. Milleporidium, eine Hydrocoralline 
aus dem Tithon von Stramberg.—Beitrage zur Paléontologie und Geologie Osterreich-Ungarns 
und des Orients 15:1-8, 2 pls. 


VOLUME 96, NUMBER 4 769 


Weissermel, W. 1913. Tabulaten und Hydrozoen.—Beitrége zur geologischen Erforschung der 
deutschen Schutzgebiete 5:84—-111, pl. 14. 


Department of Invertebrate Zoology, Smithsonian Institution, Washington, D.C. 
20560. 


PROC. BIOL. SOC. WASH. 
96(4), 1983, pp. 770-779 


ON A SMALL COLLECTION OF ENTOCYTHERID 
OSTRACODS WITH THE DESCRIPTIONS OF 
THREE NEW SPECIES 


Horton H. Hobbs, Jr. and Auden C. McClure 


Abstract.— Three new entocytherid ostracods, Ascetocythere jezerinaci, Anky- 
locythere carpenteri, and Ornithocythere thomai, infesting burrowing crayfishes, 
are described from Lee County, Virginia, Elmore County, Alabama, and Wash- 
ington County, Alabama, respectively. Reports of the ostracods infesting cray- 
fishes from nine additional collections containing burrowing crayfishes from Ken- 
tucky, Indiana, Ohio, Virginia, and West Virginia are also presented. 


This report is based upon ostracods gleaned from 12 containers in which cray- 
fishes from Alabama, Kentucky, Indiana, Ohio, Virginia, and West Virginia had 
been preserved. In each container, at least one of the crayfish (either Cambarus 
(Jugicambarus) dubius Faxon, 1884, or C. (Lacunicambarus) diogenes Girard, 
1852) that had been placed in it had been captured from a burrow. In five con- 
tainers, only representatives of one of these two burrowing species had been placed 
in the jars. Twelve species of ostracods, three of which are previously undescribed, 
were among the specimens found. Following the descriptions of the new species, 
new locality records are listed together with remarks concerning the ostracods 
and their host crayfishes. 

Previous records of the hosts of ostracods belonging to the genera Ascetocythere 
and Ornithocythere provide evidence that these ostracods are probably restricted 
to crayfishes that are recognized as primary burrowers. The same crayfishes, 
however, frequently have been found to harbor members of the other entocytherid 
genera encountered in the collections reported here. Thus we are reasonably certain 
that among those specimens from jars in which more than one host crayfish species 
was preserved, the members of Ascetocythere and Ornithocythere were symbionts 
of the burrowing crayfish; the other ostracods could have been using any or all 
of the crayfishes present as hosts. 


Ascetocythere jezerinaci, new species 
Fig. la—d 


Description.— Male: Eye pigmented, located approximately 0.28 shell length 
from anterior margin. Shell (Fig. 1a) subovate, about 1.7 times as long as high 
and margins lacking prominences and emarginations. Submarginal setae more 
abundant anteriorly and posteriorly than ventrally; none observed on dorsal mar- 
gin. 

Copulatory complex (Fig. 1b) with peniferum bearing 2 prominences extending 
anteriorly from subterminal expansion. More dorsal (anterior) process heavy, its 
length about half minimum anterior-posterior diameter of peniferum, tapering, 
and flanking anterodorsally directed penis. Ventral process shorter, more heavily 


VOLUME 96, NUMBER 4 771 


g 


Fig. 1. Ascetocythere jezerinaci (a—d) and Ankylocythere carpenteri (e-i): a, g, Dextral view of shell 
of holotype; b, c, d, e, Copulatory complex; c, d, f, Clasping apparatus; h, Clasping apparatus and 
fingers; 7, Dextral view of shell of allotype. (Scales in mm.) 


sclerotized, acute, and slightly arched. Penis complex long and extending ante- 
riorly, emerging on anterior surface of ventral expansion near tip of anterior 
process; 2 elements contiguous only along distal third of prostatic duct. Clasping 
apparatus (Fig. lc, d) arched but not clearly divisible into horizontal and vertical 
rami, massive basally, and tapering along distal four-fifths of its length; postaxial 
border entire, preaxial border with notch at base of distal third; apex with 3 or 4 
rounded denticles. Both dorsal and ventral fingers moderately robust, latter gently 
curved from base and directed anteroventrally. 

Measurements.— Holotype: length of shell 392 uw; height of shell 217 uw; corre- 
sponding measurements of the paratypic male 399 uw and 217 uy, respectively. 

Type-locality.— Creek and burrows 2.2 air kilometers (1.3 miles) north of Stick- 
leyville, Lee County, Virginia (36°33’N, 82°55’W). The specimens on which this 
description is based were removed from debris in a jar that had contained spec- 
imens of the crayfishes C. (J.) dubius, C. (Cambarus) bartonii (Fabricius, 1798), 


772 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


and C. sp. Also infesting these crayfishes were the entocytherids Donnaldsoncy- 
there donnaldsonensis (Klie, 1931) and Uncinocythere simondsi (Hobbs and Wal- 
ton, 1960). 

Disposition of types.—The holotypic and paratypic male are deposited in the 
National Museum of Natural History, Smithsonian Institution, numbers 204400 
and 204401, respectively. 

Range and specimens examined.—This ostracod is known only from the type- 
locality, from which we have seen only the two type-specimens. 

Host and entocytherid associates. —See ““Type-locality.”’ 

Relationships. — Ascetocythere jezerinaci, a member of the Asceta Group (Hobbs 
and Hart 1966:39), has its closest affinities with As. didactylata Hobbs and Hart 
(1966). Although the processes borne on the ventral extremity of the peniferum 
of the latter are directed ventrally and those of As. jezerinaci are disposed ante- 
riorly, the relative positions of the processes made them readily comparable. 
Moreover, except for fewer, only one, denticles on the preaxial border of the 
clasping apparatus in As. jezerinaci, this structure is strikingly similar in the two 
species, aS are most other features. 

Hobbs and Walton (1975:6—-7) presented a key to the then-known members of 
the genus. To include the new species described here in it, the following is offered 
to replace couplet 12 in their key. 


12(3'). Ventral part of peniferum with angular flange .................. 13 
QD, Ventral part of peniferum lacking angular flange ................ 16 
16(12'). Processes on ventral extremity of peniferum directed ventrally; 
preaxial border of clasping apparatus with more than one denticle 
RO Lot RS es MCE Ae As. didactylata Hobbs and Hart, 1966:43 
oe Processes on ventral extremity of peniferum directed anteriorly; 
preaxial border of clasping apparatus with single denticle ........ 
a tet aoa Me on Eg Sie he a ANU a Es el ae AS. jezerinaci, new species 


Etymology. —This entocytherid is named in honor of Raymond F. Jezerinac of 
Ohio State University at Newark, a student of crayfishes and one of the collectors 
of most of the material examined in this study. 


Ankylocythere carpenteri, new species 
Fig. le-i 

Description. — Male: Eye pigmented, located about 0.2 shell length from anterior 
margin. Shell (Fig. 1g) subovate, about 1.6 times as long as high, margins lacking 
prominences and emarginations. Submarginal setae present anteriorly and pos- 
teriorly, sparse ventrally, and absent dorsally. 

Copulatory complex (Fig. le) with arched peniferum moderately deeply exca- 
vate ventrally resulting in acute, anteroventrally directed prominences anteriorly 
and posteriorly, posterior prominence distinctly curved anteriorly. Penis complex 
situated in ventral sixth of peniferum but exhibiting no distinctive features. Clasp- 
ing apparatus (Fig. le, f, h) L-shaped with vertical ramus 1.6 to 1.9 times as long 
as horizontal ramus. Former slightly arched, and neither preaxial nor postaxial 
borders armed; preaxial border of horizontal ramus with single tooth near mid- 


VOLUME 96, NUMBER 4 HHS) 


length; postaxial margin with (1) distinct angular bend at or near junction of 
vertical and horizontal rami, in some views appearing rounded and slightly pro- 
duced (Fig. 1 e); (2) prominent slender talon, situated slightly proximal to level of 
tooth on preaxial border, directed anteroventrally and somewhat mesially; and 
(3) well defined excrescence slightly proximal to midway between distal base of 
talon and two apical denticles. Dorsal finger prominent, straight, and bearing 
simple apical seta; ventral finger moderately slender, curved throughout length 
or along proximal and distal fourths, and directed anteroventrally. 

Female: Eye located about 0.14 shell length from anterior margin. Shell (Fig. 
17) subovate, 1.6 times as long as high, and shallowly excavate ventrally anterior 
to midlength. Shell margin otherwise entire. Submarginal setae present but rather 
widely spaced anteriorly, ventrally, and posteriorly. 

Genital apparatus, like that of other members of genus, consisting of simple, 
slightly tapering papilla. 

Measurements.—The length of five males ranges from 378 to 399 uw, mean 
388 + 7.98 uw; the height ranges from 238 to 252 uw, mean 244 + 5.9 w; corre- 
sponding measurements of five females are 399 to 413 uw, mean 405 + 8.6 w, and 
245 to 252, mean 248 + 3.83 wu. 

Type-locality.—Small stream entering Coosa River 1.7 kilometers upstream 
from Bibb Graves Bridge at Wetumpka, Elmore County, Alabama (T.18N, R.18E, 
Sec. 13). The host was C. (L.) diogenes. 

Disposition of types.—The holotypic male, allotypic female, and a dissected 
paratypic male are deposited in the National Museum of Natural History, Smith- 
sonian Institution, numbers 204402, 204403, and 204404, respectively. Paratypic 
males are in the collections of the British Museum (Natural History) and that of 
H. H. Hobbs III, Wittenberg University. 

Range and specimens examined. — Ankylocythere carpenteri is known only from 
the type-locality where 11 specimens were obtained from five crayfish. 

Host and entocytherid associates.—See ““Type-locality.”’ 

Relationships. —Ankylocythere carpenter is allied to those members of the genus 
that have a well developed talon on the horizontal ramus of a clasping apparatus 
that does not have a conspicuously long vertical ramus. Among its closer relatives 
are Ankylocythere freyi Hobbs III, 1978; Ank. krantzi Hobbs III, 1978; Ank. 
tiphophila (Crawford, 1959); and Ank. sinuosa (Rioja, 1942). It differs from all 
other members of the genus, in possessing a distinct angle on the postaxial surface 
of the clasping apparatus at, or immediately adjacent to, the junction of the 
horizontal and vertical rami. 

Etymology.—This ostracod is named for Michael R. Carpenter of the Smith- 
sonian Institution who collected the specimens on which the above description 
is based and who has assisted one of us (HHH) in the laboratory on numerous 
occasions during the past decade. 


Ornithocythere thomai, new species 
Fig. 2 


Description.— Male: Eye pigmented, located about one-sixth shell length from 
anterior margin. Shell (Fig. 2a) subovate, 1.8 times as long as high, and greatest 
height about 1.4 times that at level of eye, margins entire; submarginal setae closer 


774 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 2. Ornithocythere thomai: a, Dextral view of shell of holotype; b, Same of allotype; c, Cop- 
ulatory complex of dissected male paratype; d—h, Clasping apparatus. (Scales in mm.) 


together anteriorly and posteriorly than ventrally and dorsally, very few along 
latter margin. 

Copulatory complex (Fig. 2c) with peniferum extending ventrally clearly beyond 
clasping apparatus, ventral part strongly sclerotized, and anterodorsally directed 
beaklike prominence with ventral subangular knob near midlength. Hyaline, tri- 
angular, lamelliform process situated at proximal base of beaklike prominence, 
its apex directed anteriorly. Penis conspicuous, U-shaped, and situated in ventral 
fifth of peniferum, its two components contiguous for about half length of prostatic 
element. Penis guides strongly sclerotized. Clasping apparatus (Fig. 2d—h) broadly 
C-shaped, not clearly divisible into vertical and horizontal rami, gently tapering 
in diameter almost from base. Preaxial border entire almost to apex, there bearing 
4 apical denticles; postaxial border also entire except for series of 4 or 5 rounded 
to subangular prominences on distal fourth. Ventral finger (excluding apical setae) 
about 4 times length of dorsal finger, straight along proximal two-thirds, distal 
third bent anteriorly at 90 degrees, bearing simple seta; dorsal finger straight, 
directed anteroventrally, and terminating in apparently bifurcate seta. 

Triunguis female: Eye situated about one-third shell length from anterior mar- 
gin. Shell (Fig. 25) about 1.7 times as long as high, truncate posteriorly, greatest 
height 1.4 times that at level of eye. Submarginal setae distributed as in male. 

Genital apparatus consisting of short, sclerotized, conical papilla; apparently 
flexible hyaline rod, coated with detritus, extending pendant from apex. 


VOLUME 96, NUMBER 4 775 


Measurements.— The length of the shells of seven males ranges from 455 to 
504 wu, mean 487 + 15.6 uw, the height from 252 to 273 uw, mean 266 + 7.0 yu; 
corresponding measurements of the two females are 504 uw and 287 to 294 un. 

Type-locality.— Burrows along drainage ditch adjacent to Crosbys Creek at Mill- 
ry on State Route 17, Washington County, Alabama (T.8N, R.3W, Sec. 25). The 
host crayfish was C. (L.) diogenes, which harbored no other entocytherids except 
Ank. freyi. 

Disposition of types.—The holotypic male, allotype, and a dissected paratypic 
male are deposited in the National Museum of Natural History, Smithsonian 
Institution, numbers 204405, 204406, and 204407, respectively. Paratypic males 
are in the British Museum (Natural History), in the collection of H. H. Hobbs 
III, Wittenberg University, and in the Smithsonian Institution. 

Range and specimens examined.— Known only from the type-locality. A total 
of eight males and four triunguis females were obtained from the six crayfish 
collected. 

Host and entocytherid associates.—See ““Type-locality.” 

Relationships. — Ornithocythere thomai has its closest affinities with O. aetodes 
Hobbs III, 1970. The two are remarkably similar in most features, particularly 
in the structure of the ventral part of the peniferum; however, the translucent 
triangular element is much more prominent in O. thomai, as is obvious since this 
structure has been overlooked previously in O. aetodes. The two species may be 
distinguished most readily by the rounded, as opposed to the angular, clasping 
apparatus of O. thomai. 

Etymology.—This ostracod is named in honor of Roger F. Thoma of the Ohio 
Environmental Protection Agency, a student of crayfishes, and the other collector 
of most of the specimens that are mentioned in this study. 


New Locality Records for Entocytherids and Their Hosts 
ALABAMA: 


1. Crosbys Creek and drainage ditch at Millry on State Route 17, Washington 
County, 22 Apr 1970; Horton H. Hobbs, Jr., collector. 

Entocytherids: Ank. freyi Hobbs, and O. thomai, new species. 

Host: C. (L.) diogenes Girard, 1852. 

Remarks.—Ankylocythere freyi was described from specimens that were in- 
festing the same host species as that reported herein from Crenshaw County, 
Alabama. Whereas the new locality falls within the range of the species recorded 
by Hobbs III, it lies slightly to the north of those records plotted by him (1978: 
Fig. 3) west of the Tombigbee River. Information relative to O. thomai is presented 
following its description above. 


2. Branch entering the Coosa River 1.7 kilometers upstream from Bibb Graves 
Bridge at Wetumpka, Elmore County, 12-13 Aug 1976; Michael R. Carpenter, 
collector. 

Entocytherid: Ank. carpenteri, new species. 

Host: C. (L.) diogenes. 

Remarks.—As noted above, this ostracod is known only from the type-locality, 
where no other entocytherids were found infesting the host. 


776 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


INDIANA: 


Roadside ditch 6.1 air kilometers west of New Point, just north of State Route 
46, Decatur County, 14 June 1980; John A. Thoma, Roger F. Thoma, and Ray- 
mond F. Jezerinac, collectors. 

Entocytherid: Dactylocythere crawfordi Hart, 1965. 

Host: C. (L.) diogenes. 

Remarks. —This is the third record of the occurrence of Dt. crawfordi in Indiana; 
the other two are in the basins of the Wabash and West Fork of the White rivers. 
The locality cited above lies in the watershed of the East Fork of the White River, 
some 85 kilometers southeast of that in Marion County listed by Hart and Hart 
(1974:55). 


KENTUCKY: 


1. Patton’s Creek (Ohio River basin), 3.4 kilometers northwest of Sligo on County 
line road, Oldham-Trimble counties, 19 Apr 1980; J.A.T., R.F.J., and Mathew 
McClusky, collectors. 

Entocytherids: Dactylocythere exoura Hart and Hart, 1966, Dt. ungulata (Hart 
and Hobbs, 1961), and Donnaldsoncythere donnaldsonensis (Klie, 1931). 

Hosts: Cambarus (C.) ortmanni (Williamson, 1907), C. (Erebicambarus) or- 
natus Rhoades, 1944a, C. (L.) diogenes, and Orconectes rusticus (Girard, 1852). 

Remarks. — Hart and Hart (1974:58) recorded only two localities for Dt. exoura, 
one each in Grant and Oldham counties. The new locality is no more than six or 
seven kilometers north of that in Oldham County cited by the Harts, and the 
hosts are among those previously cited. Although the Oldham-Trimble locality 
does not alter the limits of the range of Dt. ungulata depicted by Hart and Hart 
(Fig. 49), it does provide a new county record and a precise locality on the 
northwestern boundary. Neither this nor any of the following records for the wide- 
ranging Dn. donnaldsonensis is noteworthy. 


2. Big Creek (tributary to Levisa Fork, Big Sandy River basin), 3.4 kilometers 
northeast of Dunlap on State Route 194, Pike County, 18 Jun 1981; R.F-.T., 
Raymond J. Jezerinac, and R.F.J., collectors. 

Entocytherids: Ascetocythere sclera Hobbs and Hart (1966) and Dn. donnald- 
sonensis. 

Hosts: Cambarus (C.) sciotensis Rhoades, 1944b, C. (J.) dubius, Cambarus sp., 
and Orconectes sp. 

Remarks.—To our knowledge, this is the first record of the occurrence of As. 
sclera in Kentucky; however, having been found in neighboring parts of Virginia 
and West Virginia (Hart and Hart 1974:41), its occurrence in the extreme south- 
eastern part of Kentucky is not a surprise. Our specimens exhibit no variations 
that set them apart from those from elsewhere. 


OHIO: 


Vernal pond, 6.9 kilometers northeast of Hebron on Licking Twp. Road 305, 
Licking County, 9 Apr 1981; R.F.T. and Craig Ciola, collectors. 

Entocytherid: Dt. crawfordi. 

Host: C. (L.) diogenes. 


VOLUME 96, NUMBER 4 VHT 


Remarks.—This record extends the known range of Dt. crawfordi about 60 
kilometers to the east into the Muskingum River basin. 


VIRGINIA: 


1. Along Hobbs Branch (tributary of Levisa Fork, Big Sandy River basin), about 
1.0 kilometer southeast of State Route 35, 5.1 kilometers east of Grundy, Bu- 
chanan County, 18 Jun 1981; R.F.T., R.F.J., and R.J.J., collectors. 
Entocytherids: As. sclera and Dn. donnaldsonensis. 
Hosts: C. (C.) sciotensis, C. (J.) dubius, and Cambarus sp. 


2. 2.2 air kilometers north of Stickleyville, along tributary to Wallen Creek, Powell 
River basin, Lee County, 20 Jun 1981; R.F.J. and R.F.T., collectors. 
Entocytherids: As. jezerinaci, new species, Dn. donnaldsonensis, and Uncino- 
cythere simondsi. 
Hosts: C. (C.) cavatus Hay, 1902, C. (J.) dubius, and Cambarus (C.) sp. 
Remarks.— All of the information available to us concerning As. jezerinaci is 
presented immediately following the description of this entocytherid. The presence 
of U. simondsi here represents a new state record. 


3. Spring seep along State Route 871, 0.7 kilometer north of the junction with 
Route 646, Scott County, 9 Jul 1981; collectors (?). 

Entocytherids: Ascetocythere ozalea Hobbs and Hart, 1966; and Dn. donnald- 
SonensIs. 

Host: Cambarus (J.) dubius. 

Remarks.—This 1s the third locality reported for As. ozalea which, insofar as 

we are aware, 1s endemic in the Tennessee River basin in Virginia; the other two 
localities are in Scott and Russell counties, Virginia (Hart and Hart 1974:40). 
4. Mud Fork (Bluestone-New River basin), about 0.9 kilometer north of the town 
of Mud Fork (17.9 kilometers west southwest of Bluefield, West Virginia), on 
County Road 643, 19 Jun 1981; R.F.T., R.F.J., and R.J.J., collectors. 

Entocytherid: Dn. donnaldsonensis. 

Hosts: C. (C.) sciotensis, C. (J.) dubius, and Cambarus. sp. 


WEST VIRGINIA: 


1. Panther Creek State Forest (Tug Fork-Big Sandy River basin), 10 kilometers 
south of Panther, McDowell County, 19 Jun 1981; R.F.T., R.F.J., and R.J.J., 
collectors. 

Entocytherids: Du. donnaldsonensis, Phymocythere phyma (Hobbs and Walton, 
1962), and U. simondsi. 

Hosts: Cambarus (C.) sciotensis, C. (J.) dubius, Cambarus sp., and Orconec- 
tes sp. 

Remarks.—The presence of P. phyma at this locality represents the south- 
westernmost record for the species, also the first report of its presence in the 
Guyandot drainage system. Uncinocythere simondsi has not been previously re- 
ported from West Virginia. 

2. Tributary of West Fork of Twelve Pole Creek (Ohio River basin), Cabwaylingo 
State Forest, 4.3 kilometers northwest of Wilsondale, Wayne County, 18 Jun 
1981; R.F.T., R.F.J., and R.J.J., collectors. 


778 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Entocytherids: As. sclera and Donnaldsoncythere donnaldsonensis. 
Hosts: C. (J.) dubius and Cambarus sp. 


Acknowledgments 


We are grateful to Michael R. Carpenter, Raymond F. Jezerinac, and to Roger 
F. Thoma for donating most of the specimens reported here to the Smithsonian 
Institution, and to Messrs. Jezerinac and Thoma for supplying us with the iden- 
tifications of the hosts of the entocytherids collected by them and for their com- 
ments on the manuscript. Thanks are also extended to C. W. Hart, Jr., of the 
Smithsonian Institution and to H. H. Hobbs III of Wittenberg University for 
their criticisms of the manuscript, and to Margaret Webb of the Madeira School, 
Greenway, Virginia, for assistance in preparing the illustrations. 


Literature Cited 


Crawford, E. A. 1959. Five new ostracods of the genus Entocythere (Ostracoda, Cytheridae) from 
South Carolina.— University of South Carolina Publications, Series III, Biology, 2(4):149-189, 
37 figures. 

Fabricius, Johann C. 1798. Supplementum Entomologiae Systematicae, 572 pages. Hafniae et Storch. 

Faxon, Walter. 1884. Descriptions of new species of Cambarus, to which is added a synonymical 
list of the known species of Cambarus and Astacus.— Proceedings of the American Academy 
of Arts and Sciences 20:107-158. 

Girard, Charles. 1852. A revision of the North American Astaci, with observations on their habits 
and geographical distribution. — Proceedings of the Academy of Natural Sciences of Philadelphia 
6:87-91. 

Hart, C. W., Jr. 1965. New entocytherid ostracods and distribution records for five midwestern 

states. —Transactions of the American Microscopical Society 84(2):255—259, 5 figures. 

, and Dabney G. Hart. 1966. Four new entocytherid ostracods from Kentucky, with notes on 

the troglobitic Sagittocythere barri.—Notulae Naturae of the Academy of Natural Sciences of 

Philadelphia 388:1—10. 

, and Horton H. Hobbs, Jr. 1961. Eight new troglobitic ostracods of the genus Entocythere 

(Crustacea, Ostracoda) from the eastern United States. — Proceedings of the Academy of Natural 

Sciences of Philadelphia 113(8):173-185, 32 figures. 

Hart, Dabney G., and C. W. Hart, Jr. 1974. The ostracod family Entocytheridae.—Academy of 
Natural Sciences of Philadelphia Monograph 18:ix + 239 pages, 49 figures, 52 plates. 

Hay, William Perry. 1902. Observations on the crustacean fauna of Nickajack Cave, Tennessee, and 
vicinity. — Proceedings of the United States National Museum 25(1292):417—439, 8 figures. 

Hobbs, Horton H.., Jr., and C. W. Hart, Jr. 1966. On the entocytherid ostracod genera Ascetocythere, 

Plectocythere, Phymocythere (gen. nov.), and Cymocythere with descriptions of new species. — 

Proceedings of the Academy of Natural Sciences of Philadelphia 118(2):35—-61, 37 figures. 

,and Margaret Walton. 1960. Three new ostracods of the genus Entocythere from the Hiwassee 

drainage system in Georgia and Tennessee.—Journal of the Tennessee Academy of Science 

35(1):17-23, 20 figures. 

, and 1962. New ostracods of the genus Entocythere from the Mountain Lake region, 

Virginia (Ostracoda, Entocytheridae).— Virginia Journal of Science 13(2):42—48, 13 figures. 

, and 1975. New entocytherid ostracods from Tennessee with a key to the species of 

the genus Ascetocythere.— Proceedings of the Biological Society of Washington 88(2):5—20, 2 

figures. 

Hobbs III, H. H. 1970. New entocytherid ostracods of the genus Ornithocythere and the description 

of a new genus. — Proceedings of the Biological Society of Washington 83(15):171—182, 2 figures. 

1978. New species of ostracods from the Gulf Coastal Plain (Ostracoda: Entocytheridae).— 
Transactions of the American Microscopical Society 97(4):502—511, 6 figures. 
Klie, W. 1931. Campagne spéologique de C. Bolivar et R. Jeannel dans |’Amérique du Nord (1928), 


VOLUME 96, NUMBER 4 VS) 


3:Crustaces Ostracodes. — Biospeologica: Archives de Zoologie Expérimentale et Générale 7 1(3): 
333-344, 20 figures. 

Rhoades, Rendell. 1944a. The crayfishes of Kentucky, with notes on variation, distribution and 

descriptions of new species and subspecies.— American Midland Naturalist 31(1):111-149, 10 

figures. 

. 1944b. Further studies on distribution and taxonomy of Ohio crayfishes and the description 

of a new subspecies.— Ohio Journal of Science 44(2):95-99. 

Rioja, Enrique. 1942. Estudios carcinologicos. XIV, Consideraciones y datos acerca del género 
Entocythere (Crus. Ostracodos) y algunas de sus especies, con descripcion de una nueva.— 
Anales del Instituto de Biologia, Universidad Autonoma de Mexico 13(2):685-697, 21 figures. 

Williamson, E. B. 1907. Notes on the crayfish of Wells County, Indiana, with description of new 
species. — 31st Annual Report of the Department of Geology and Natural Resources, Indiana 
1906:749-763, plate 35. 


(HHH) Department of Invertebrate Zoology, Smithsonian Institution, Wash- 
ington, D.C. 20560; (ACM) 823 Bulls Neck Road, McLean, Virginia 22102. 


PROC. BIOL. SOC. WASH. 
96(4), 1983, pp. 780-806 


SEVEN NEW SPECIES OF THE INDO-PACIFIC 
GENUS EVIOTA (PISCES: GOBIIDAE) 


Susan L. Jewett! and Ernest A. Lachner 


Abstract.— Seven new species of gobiid fishes of the genus Eviota Jenkins from 
Indo-Pacific marine waters are described as: E. albolineata (a member of Group 
I, Lachner and Karnella 1980:113); E. japonica, E. latifasciata, E. punctulata 
(Group II); E. cometa, E. sigillata (Group II); and E. sparsa (Group VII, the 
characters of which are discussed herein). The available evidence does not indicate 
that these species form a natural group. A table of characters for Groups I, I, II, 
and VII is given. Illustrations of the seven new species and a table of pertinent 
characters and meristics are provided. 


As part of our continuing study of the systematics and zoogeography of the 
gobiud genus Eviota Jenkins (Lachner and Karnella 1978, 1980; Karnella and 
Lachner 1981), we present descriptions of seven new species. In our earlier study 
(1980:113) we discussed six species groups of Eviota. We herein define a seventh 
species group, and relate each of the new species to Groups I, II, III, or VII. A 
summary of species group characters is presented in Table 1. 

The new species and their species group allocations are as follows: Eviota al- 
bolineata, Group I (containing 17 nominal species); E. japonica, E. latifasciata 
and E. punctulata, Group II (11 nominal species); E. cometa and E. sigillata, 
Group III (7 nominal species); and E. sparsa, Group VII (2 species, one to be 
described subsequently). The salient characters of the seven new species are given 
in the diagnostic accounts under each species, and some are summarized in 
Table 2. 


Methods 


The methods of obtaining counts and measurements and the presentation of 
these data, as well as the description of the cephalic sensory pore and cutaneous 
papillae systems, follow that of Lachner and Karnella (1978, 1980), and Karnella 
and Lachner (1981) with the following modifications: 

1. “Pelvic fin membrane” refers to the membrane connecting the third and 
fourth rays of the pelvic fin. It is considered to be reduced when its length does 
not extend to the first branch of the fourth pelvic fin ray and to be well developed 
when it exceeds that point (Fig. 1, Lachner and Karnella 1980). 

2. ““Postanal midline spots” refers to the dark spots, composed at least in part 
of subcutaneous pigmentation, that occur along the posteroventral midline of the 
trunk. These spots begin at, or just posterior to, the origin of the anal fin, and 
extend to a vertical drawn 2 to 3 scale rows anterior to the hypural joint, otherwise 
referred to as the ““midcaudal peduncle.”’ There are usually 4 to 6 postanal midline 


' Formerly Susan J. Karnella. 


VOLUME 96, NUMBER 4 781 


Table 1.—Characters distinguishing four species groups of Eviota. 
ee 


Groups 
Characters I II Ii VII 
ee ea eee 
Cephalic sensory pore 1 2 2 5 
pattern 
Vertebrae 26 26 25 26 
Some pectoral fin yes! yes no yes 
rays branched 
Male genital papilla nonfimbriate nonfimbriate, nonfimbriate nonfimbriate 
fimbriate or 
cup-shaped 
Pelvic fin membrane reduced to well reduced reduced to well _ welll developed 
developed developed 
Length of fifth pelvic absent to 2/10 absent to 1/10 1/10 to 5/10 6/10-9/10 
fin ray relative to 
fourth fin ray 
Spinous dorsal fin Yes or no yes or no yes yes 


elongation 


' Except some specimens of E. monostigma and E. pseudostigma. 


spots. An additional spot, usually smaller and less intense, may occur more pos- 
teriorly, near the insertion of the ventral procurrent rays. This spot is not included 
when counting the postanal midline spots. 

3. ““Midcaudal peduncle spot’’ refers to a dark spot on the caudal peduncle, 
usually centered on the lateral midline, 2 to 3 scale rows anterior to the insertion 
of the caudal fin. The spot may be composed entirely of subcutaneous pigment 
or may be a combination of surface and subcutaneous pigmentation. It is in line 
with the last subcutaneous trunk bar when a series of trunk bars occurs. 

Three cephalic sensory pore patterns are found in the species described in this 
paper. Pore pattern | represents the full complement of sensory pores for the 
genus Eviota and includes the paired nasal (NA) pores, the single anterior inter- 
orbital (AITO) and posterior interorbital (PITO) pores, the paired supraotic (SOT), 
anterior otic (AOT), and intertemporal (IT) pores, as well as two pairs, an upper 
and lower, of preopercular (POP) pores (Fig. 4, Lachner and Karnella 1980). Pore 
pattern 2 lacks only the IT pores, and pore pattern 5 lacks the IT and both pairs 
of POP pores. Cutaneous papillae patterns A, B and B-1 (Lachner and Karnella 
1980:7) correspond to pore patterns 1, 2 and 5, respectively. 

Presentation of material examined for holotypes and paratypes is of the fol- 
lowing format: catalog number, size range, abbreviated locality data, depth of 
capture, collector and field number. Non-type material is summarized by geo- 
graphic locality rather than listed by individual museum lot. 

Abbreviations: the following museum acronyms are used to designate institu- 
tions and collections cited: 


AMNH American Museum of Natural History, New York 
AMS Australian Museum, Sydney 

ANSP Academy of Natural Sciences, Philadelphia 
BPBM _ Bernice P. Bishop Museum, Honolulu 


PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


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VOLUME 96, NUMBER 4 783 


CAS California Academy of Sciences, San Francisco 

FMNH_ Field Museum of Natural History, Chicago 

ROM Royal Ontario Museum, Toronto, Canada 

RUSI Rhodes University, J.L.B. Smith Institute of Ichthyology, Grahams- 
town, South Africa 

USNM_ Former United States National Museum, now National Museum of 
Natural History (NMNH), Smithsonian Institution, Washington, D.C. 

WAM Western Australian Museum, Perth 

YCM Yokosuka City Museum, Yokosuka, Japan 


Eviota albolineata, new species 
Figs. 1-2 


Material examined.—998 specimens from numerous localities, ranging from 
the east coast of Africa to the Tuamotu Archipelago; total size range 7.7—24.7; 
gravid females 11.1—21.1. 

Holotype: USNM 227140, (22.2), male; Tahiti, shallow patch south of Tap- 
ueraha Pass, 0-3.0 m, 21 Apr 1970, C. L. Smith, S70-45. 

Paratypes: TAHITI: AMNH 43023, 2 (16.2, 21.3); same data as holotype. 
AMNH 43022, 1 (16.4); north side of Passe Tiamahana, 10.7-13.7 m, S70-11. 
AMNH 43025, 1 (18.9), off Papeari, O-2.1 m, S70-53. AMNH 43024, 13 (15.6- 
19.3); south of Tapueraha reef, 0-3.6 m, S70-41. USNM 227166, 11 (14.1-—22.0); 
south of Tapueraha, 0-7.6 m, S70-44. CAS 52829, 4 (16.4—21.6):; off Papeari, 
6.1-10.7 m, S70-51. BPBM 29191, 1 (19.4); off Papeari, 0-3.6 m, S70-55. AMS 
1.24025-001, 6 (16.0-19.4); south of Tapueraha Pass, 0—2.4 m, S70-43. ANSP 
151994, 6 (16.9-19.8); same data as previous. CAS 48471, 4 (14.4-19.5); Ati- 
maono, Teauaraa Pass, 0.9-13.7 m, sta 18, GVF Reg. 1350. HUAHINI NUI: 
AMNH 43034, 45 (13.2-20.9); 0.5 mi. south of Fare, 10.7-12.2 m, S70-8. The 
following five lots with same data as previous: USNM 227165, 5 (15.4-19.9); 
ANSP 151995, 5 (13.5-22.0); BPBM 29192, 5 (14.6—20.6); CAS 52830, 5 (13.7- 
21.1); AMS 1.24026-001, 5 (13.2—22.9). AMNH 43033, 1 (17.9); 0.5 mi. south 
of Passe Avapeihi, 0-1.8 m, S70-6. AMNH 43020, 2 (14.4, 19.9): ca. 2 mi. south 
of Fare, 0-1.8 m, S70-7. BORA BORA: AMNH 43027, 5 (12.7-19.0); lagoon 
channel south of Topua Is., 0O-7.6 m, S70-19. AMNH 43028, 1 (13.5); ca. 2 mi. 
southwest of Topua Is., O-1.8 m, S70-16. AMNH 43026, 1 (19.7); 2 mi. southwest 
of Topua Is., 0O-1.8 m, S70-15. USNM 227167, 7 (13.5-19.3); lagoon channel 
south of Topua Is., 0O—-10.7 m, S70-18. 

Non-type Material: Numerous specimens from the following localities: 
OCEANIA: Society Islands, Tuamotu Archipelago, Tubuai Islands, Cook Islands, 
Samoa Islands, Fiji, New Hebrides, Santa Cruz Islands, Solomon Islands, Gilbert 
Islands, Marshall Islands, Marianas Islands, Caroline Islands, Palau Islands; PA- 
PUA NEW GUINEA; AUSTRALIA: Lord Howe Island, Great Barrier Reef; 
TAIWAN; PHILIPPINE ISLANDS; INDONESIA; INDIAN OCEAN: India, Sri 
Lanka, Chagos Archipelago, Agalega Islands, Mauritius, Seychelles Islands, Ami- 
rantes Islands, Aldabra Atoll, Comoro Islands, Mozambique. 

Diagnosis.— Cephalic sensory pore system complete; pectoral fin rays numerous, 
modally 18, with most rays branched; dorsal/anal fin ray formula typically 9/8; 
elongation of spines in first dorsal fin uncommon; pelvic fin typically I, 4 1/10- 


784 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 1. Eviota albolineata. a, 16.4 mm SL, male, Seychelles Islands, USNM 227132; b, 20.3 mm 
SL, male, Tuamotu Archipelago, BPBM 14048; c, Holotype, 22.2 mm SL, male, Tahiti, USNM 
227140. 


2/10 with well developed membrane between rays; branches on fourth pelvic fin 
ray 6-12; trunk lacking both dark spot on middle portion of caudal peduncle and 
well developed postanal midline spots although some tiny spots or dark streaks 
may be present; body generally pale with dark dusky anal fin and scattered spec- 
kling on second dorsal and caudal fins; when color pattern highly developed, as 
in eastern Oceania populations, head with weak alternating dark and light hori- 
zontal stripes dorsolaterally and trunk moderately to distinctly dusky. 

Description.— Dorsal fin VI-I, 8(1), VI-I, 9(20); anal fin I, 8(27); pectoral fin 
rays 16(2), 17(5), 18(7), 19(6), 20(1); pectoral fin rays 4-18 may be branched, 7— 
17 usually branched; pelvic fin I, 4 plus a rudiment (1), I, 4 1/10(/6), I, 4 2/10(4); 
branches on fourth pelvic fin ray 6-12, average 8.4; segments between consecutive 
branches of fourth pelvic fin ray 1—4, average 2.1; pelvic fin membrane well 
developed; branched caudal fin rays 11(8), 12(8), 13(2), 14(1); segmented caudal 
fin rays 16(1), 17(20); lateral scale rows 23(4), 24(13), 25(2); transverse scale rows 
6(2), 7(11); breast scaleless; vertebrae 10(13) precaudal plus 16(/2), 17(1) caudal, 
total 26(/2), 27(1). 

Dorsal fin elongation uncommon, observed only in males, usually involving 


VOLUME 96, NUMBER 4 785 


fe 
© BE Aus Bie 


Fig. 2. Eviota albolineata, 22.2 mm SL, male, Moorea, Society Islands, CAS 48469. 


first spine, occasionally second spine; maximum length of depressed first spine 
extending to end of base of second dorsal fin. Pelvic fin usually not extending 
beyond origin of anal fin. Cephalic sensory pore system pattern 1; cutaneous 
papillae system pattern A. Male genital papilla not fimbriate. 

Slender species, trunk beneath spinous dorsal fin not very deep relative to depth 
of head and caudal peduncle. Distal margin of nonfilamentous spinous dorsal fin 
straight or slightly concave and approximates a vertical line perpendicular to 
horizontal axis of body, in contrast to a straight margin forming acute angle with 
horizontal axis, as in most other Eviota. 

Color of preserved specimens.— This species is a complex of two color forms, 
one highly pigmented, represented in part by the type material from the Society 
Islands and Tuamotu Archipelago (eastern Oceania population) and a pallid form 
that is more widespread, occurring more to the west in Oceania, the Indo-Aus- 
tralian Archipelago, the Great Barrier Reef, and the Indian Ocean (western Ocea- 
nia-Indian Ocean population). Within the geographic ranges of each color form 
are irregularly distributed intermediate color forms which are not typically clinal. 

The coloration typical of the highly pigmented specimens of eastern Oceania 
(see Figs. 1c, 2) follows. Dorsolateral portion of head behind eyes with 2 or 3 
narrow dark horizontal stripes composed of moderately dense scattered brown 
chromatophores, upper 2 stripes extending to anterior section of trunk, lower 
stripe extending to anterior portion of opercle. Lowermost dark stripe commonly 
obscure, other 2 varying in intensity and length. Dark stripes separated by 3 pale 
bands equal to or wider than stripes, upper pale band most prominent but some- 
times masked by pigmentation on dorsal portion of head and nape. Pale bands 
often indistinct, uppermost may be limited to a large pale spot dorsolaterally 
behind eye at about position of supraotic pore. Striped pattern variously reduced 
or absent in some specimens. Head and nape dorsally variably pigmented, either 
with chromatophores scattered uniformly over predorsum, with weak to mod- 
erately developed saddles traversing midline, or with reticulated pattern; predorsal 
midline with 2 elongate spots evenly spaced between eyes and dorsal fin origin; 
head laterally with 3 small dusky spots, | at upper preopercular pore, 2 bordering 
margin of eye at four o’clock and two o’clock, latter at about anterior otic pore; 


786 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


3 spots most apparent when dark horizontal stripes faint or absent; spot at upper 
preopercular pore most persistent of 3. Central portion of chin pale or rarely with 
small discrete spot in line with vertical through middle of eye, or with 2 to 4 
small spots lateral to this position. Tip of snout and region around anterior nasal 
tubes dusky. Remainder of head mostly pale. 

Fleshy base of pectoral fin variously pigmented with scattered chromatophores: 
most commonly, upper third with pale round to horizontally elongate spot more 
or less surrounded by faint chromatophores, or, uncommonly, with faint scattered 
chromatophores throughout or pale spots on both upper and lower portions of 
base. 

Trunk often with rather uniformly scattered brown chromatophores, somewhat 
more dense dorsolaterally, in some specimens chromatophores absent ventrally. 
Dorsal portion of trunk, especially along insertions of dorsal fins, may have small 
clusters of chromatophores. Lateral midline of trunk with pale horizontal stripe 
about a half scale in width, stripe often obscure or lacking; rarely, a weak pale 
horizontal stripe along middorsolateral portion of trunk. 

Tiny dark spots usually present along ventral midline of trunk posterior to 
origin of anal fin; spots number up to 14, not integrated with any subcutaneous 
pigment; spots sometimes joined to form variously developed elongate streaks. 
Subcutaneous postanal midline spots and bars and midlateral caudal peduncle 
spot typical of many Eviota absent in this species. However, subcutaneous pig- 
mentation present in belly region as | to 3 large dark patches. 

First dorsal fin pale to dusky, usually with some dusky pigmentation posteriorly 
on distal portion of membrane, and sometimes with narrow dark basal band; in 
some specimens, fin dusky throughout. Second dorsal and caudal fins with discrete, 
fine, scattered, brown chromatophores on otherwise pale membrane; in some very 
darkly pigmented specimens (including holotype) second dorsal and caudal fins 
dusky brown throughout, lacking distinct dark spots, but with discrete small pale 
spots interspersed over membrane. Anal fin uniformly very dark brown. Pectoral 
and pelvic fins usually pale except on very dark specimens where finely dusky. 

Coloration of widespread pallid form (see Fig. 1a), found primarily in Indian 
Ocean and western Oceania, consisting of very pale head and trunk, with traces 
of most of diagnostic marks listed above for eastern Oceania form. Remnants of 
light and dark horizontal stripes on head may persist; upper pale band may be 
represented by pale spot near the supraotic sensory pore and lower dark stripe by 
narrow band of chromatophores. Three small dark spots at upper preopercular 
sensory pore and at the margin of the eye at two o’clock and four o’clock, usually 
persist. Chin usually with small dark spot on middle portion, nearly in line with 
vertical through eye, sometimes replaced or supplemented by 2 to 4 dark lateral 
spots. Predorsal midline with scattered chromatophores anteriorly, 1, or less fre- 
quently 2, elongate dark streaks mesially and weak remnants of transverse bar 
posteriorly. Fleshy base of pectoral fin most often pale, sometimes with weak 
scattered chromatophores in midsection. Trunk usually entirely pale, at most with 
scattered brown chromatophores on upper portion; sometimes pale midlateral 
band faintly discernible. Ventral midline of posterior trunk with tiny dark spots 
or continuous streak, as in dark color form, but intensity of pigmentation some- 
what reduced; some specimens have this pigmentation segmented into 6 or fewer 


VOLUME 96, NUMBER 4 787 


elongate dark spots, but pigment never as extensive as in species of Eviota with 
well developed postanal midline spots. 

First dorsal fin largely pale, distal portion with some faint scattered speckling 
and some specimens with weak narrow dusky basal band. Second dorsal fin pale 
in anterior basal region, remainder of fin pale to light dusky with fine dark spots 
scattered over midportion. Caudal fin pale to dusky with fine dark spots scattered 
over upper two-thirds of fin. Anal fin in males moderate to dark dusky throughout; 
in females fin may be somewhat lighter and pigmentation restricted to distal or 
middle portion of fin or, occasionally, lacking. Pectoral and pelvic fins pale. 

The heavily pigmented color form, with well developed dark and light stripes 
on head, is found in the Society Islands and Tuamotu Archipelago, and is also 
represented by a few specimens from the Cook and Santa Cruz Islands, and the 
Great Barrier Reef. Specimens with only traces of this pattern are found in a wider 
geographic area, but occur irregularly in a non-clinal distribution. Specimens from 
all localities in the Indian Ocean other than Sri Lanka, and most specimens from 
Australia show no traces of the head stripes. When the dark stripes are not 
discernible the species may be recognized by the characteristic dark spots on the 
side of the head, by the dark anal fin, the characteristically spotted second dorsal 
and caudal fins, the lack of subcutaneous ventral midline trunk spots and bars, 
and the generally pale trunk. The single dark spot on the chin is more common 
in specimens from the Indian Ocean than in those from western Oceania, and is 
almost entirely lacking in eastern Oceania. 

Color in life.—Indian Ocean color form. The following observations were made 
by Thomas H. Fraser at Inhaca Island, Mozambique (RUSI 1856): Specimen 
taken from hole in rocky substrate at base of coral head; “‘brilliant solid green, 
very conspicuous,” observed under water at distances of about 4.5—6 m. 

Western Oceania color forms. The following color notes were recorded by R. 
Wass. Specimens taken at Tutuila Island, Samoa Islands (USNM 222522): “Body 
pale, edges of scales orange, yellow spot on pectoral base, yellow spots on anal 
base, dusky distally, orange spots on head and lips, seven orange internal blotches 
behind anus.” (USNM 222520): ““Seven internal orange marks behind anus, body 
orange especially dorsally, nape with four orange saddles, lips orange, snout yellow, 
upper pectoral base red, trailing edges of fins dusky.” The following color was 
described for a specimen from Guam (UG 4324): ““when live—transparent with 
pink and green patches along back and lower sides.” 

Geographic distribution.—A widely distributed and abundant species, ranging 
from the east coast of Africa eastward through the Indo-Pacific region to the 
Tuamotu Archipelago. This species is not known from the Red Sea, Japan, and 
the Hawaiian Islands. 

Etymology.—The specific name is a Latin combination meaning white line, in 
reference to the pale stripes situated laterally on the head. 

Remarks.—The existence of two color forms of Eviota albolineata presents a 
systematic problem that is further compounded by the close relationship of E. 
albolineata with the allopatric species E. guttata. Our recognition of E. albolineata 
as a wide ranging species distinct from FE. guttata of the Red Sea and Gulf of 
Oman, is based on the examination of hundreds of specimens and the appraisal 
of several specific color marks. Yet, we regard our interpretation of these species 


788 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


as tentative because of the great differences between the pallid and the heavily 
pigmented forms of E. albolineata, and the close relationship between E. guttata 
and E. albolineata. A summary of the basis for our decision follows. 

Eviota albolineata is a member of Group I (Lachner and Karnella 1980:113), 
closely resembling E. guttata in general color pattern and meristic characters. In 
addition to Group I characters these two species share the following: three spots 
laterally on the head, at two and four o’clock behind the eye, and one at the upper 
preopercular pore; a uniformly dark anal fin; small, discrete dark spots on the 
second dorsal and caudal fins; one or more spots on the chin; and varying amounts 
of speckled pigmentation on the trunk. 

Eviota albolineata differs primarily from E. guttata in lacking the 6—7 enlarged, 
dark ventral midline spots and associated dark subcutaneous marks on the pos- 
terior trunk. Other differences are the pale body of the western Oceania-Indian 
Ocean form, the head stripes of the eastern Oceania form and the lack of a well 
developed dark bar along base of the spinous dorsal fin in both color forms of E. 
albolineata. There are minor differences in the pectoral fin ray counts: EF. guttata, 
average 16.6 range (15-18); E. albolineata, Indian Ocean, 17.1 (16-18); eastern 
Oceania, 18.7 (18-20). 

The data reported by us for color in life for the Indian Ocean color form of E. 
albolineata (Mozambique) and the western Oceania color forms (Samoa and Guam) 
are strikingly different, adding to the confusion in interpreting this species. 


Eviota japonica, new species 
Figs. 3-4 


Material examined.—67 specimens from several localities in Japan and the 
Ryukyu Islands; total size range 10.6—24.1; gravid females 13.9-17.7. 

Holotype: USNM 221758, (17.5), female; Ryukyu Is., Kohamajima Is., 9 Apr 
1974, M. Hayashi and T. Itoh, sta 9, formerly YCM-P1420. 

Paratypes: RYUKYU ISLANDS: YCM-P1459, 1 (17.7); Ishigakijima Is., Ka- 
bira Bay, M. Hayashi and T. Itoh. YCM-P2841, 4 (10.9-13.9); Ishigakijima Is., 
Kabira Bay, M. Hayashi and T. Itoh. YCM-P2615, | (15.0); Ishigakijima Is., 
Shitafukigawa River, M. Hayashi and T. Itoh. USNM 221752, 6 (10.6-16.8); 
N’afa Okinawa, Luchu Is., A/batross. JAPAN: USNM 221745, 26 (12.1-21.3); 
Tanegashima Is., Albatross. USNM 221748, 28 (11.4—24.1); Tanegashima Is., 
Albatross. FANH 94179, 19 (9.6—20.9); Aikawa Rikuzen, A/batross. 

Diagnosis.—Cephalic sensory pore system lacks IT pore; pectoral fin with some 
branched rays; dorsal/anal fin ray formula typically 9/8; spinous dorsal fin may 
contain filamentous spines in both sexes; pelvic fin I, 4 1/10—2/10 and with reduced 
membrane between rays; branches on fourth pelvic fin ray 8-16; trunk with 6 
dark postanal midline spots contiguous with subcutaneous bars, last of which 
aligned with dark midcaudal peduncle spot; 4 narrow dark bars on belly; 2 prom- 
inent dark occipital spots in addition to many smaller and less intense spots 
elsewhere on head and fleshy base of pectoral fin. 

Description.— Dorsal fin VI-I, 8(1), VI-I, 9(/8), VI-I, 10(1); anal fin I, 8(20); 
pectoral fin 15(1), 16(/4), 17(5); pectoral fin rays 8-17 may be branched, 11-16 
usually branched; pelvic fin I, 4 1/10(/9), I, 4 2/10(10); branches on fourth ray 


VOLUME 96, NUMBER 4 789 


Fig. 3. Eviota japonica. a, b, Paratype, 21.0 mm SL, female, Aikawa Rikuzen, Japan, FMNH 
94179; c, Holotype, 17.5 mm SL, female, Ryukyu Islands, USNM 221758. 


of pelvic fin 8-16, average 11.2; segments between consecutive branches of the 
fourth pelvic fin ray 0-5, average 1.4; pelvic fin membrane reduced: branched 
caudal fin rays 13(5), 14(6), 15(4); segmented caudal fin rays 17(20); lateral scale 
rows 23(4), 24(9), 25(6); transverse scale rows 7(8), 8(7); breast scaleless; vertebrae 
10(/6) precaudal and 16(/6) caudal, total 26. 

First and second spines of spinous dorsal fin may be filamentous in both sexes, 
first spine longest and may extend to end of base of second dorsal fin when 
depressed. Pelvic fin variable in length, most often not extending to origin of anal 
fin, sometimes beyond. Cephalic sensory pore system pattern 2; cutaneous papillae 
system pattern B. Male genital papilla not fimbriate. 

Color in preserved specimens. —Salient color pattern consisting of 2 prominent 
dark occipital spots, less intense dark spots and bars on head laterally and ven- 
trally, 2 dark spots on fleshy base of pectoral fin, numerous small dark spots along 
dorsal midline, and 6 postanal ventral midline spots. 

Head with 2 small dark spots on either side of dorsal midline behind eyes, 
followed by 2 pairs of prominent dorsolateral occipital spots; first pair almost 
always irregular in shape and vertically elongate, often bridged dorsally by narrow 
band of less dense chromatophores; second pair of occipital spots larger than first, 
more regular in shape, roundish to angular or triangular, and never connected to 
each other dorsally although sometimes a small dark spot or spots on dorsal 
midline midway between occipital spots; both pairs of occipital spots usually 
dense, dark brown, at least in lower portion of spot, dorsal portion, or in some 
cases whole spot, consisting of large dark chromatophores not coalesced into solid 


790 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 4. Eviota japonica, Paratype, 18.3 mm SL, female, Tanegashima, Japan, USNM 221745. 


dark pigmentation; 3 weak spots or narrow, poorly defined, transverse bars oc- 
curring along dorsal midline posterior to second pair of occipital spots; remainder 
of nape pale. 

Cheek and opercle with irregularly shaped spots and bars, those on cheek less 
intense than those on opercle, but neither as dark as occipital spots; consistently 
present pigmentation includes elongate bar below the eye at six o’clock to below 
rictus, mark from eye at about seven-thirty, across lips to chin; snout weakly 
pigmented with small spots; lower opercle, branchiostegal membranes and lower 
portion of head with many small dark irregularly shaped elongate spots, equal in 
intensity or darker than opercle spots, sometimes approaching intensity of occip- 
ital spots. 

Fleshy base of pectoral fin with 2 dark oval or elongate spots composed of loose 
aggregates of large dark chromatophores, spots clearly separated in midportion 
of base, both spots about equal in intensity to opercle spots. Dorsal midline with 
approximately 13-14 small dark spots or narrow bars from origin of spinous 
dorsal fin to procurrent rays of caudal fin. Six small, dark postanal midline spots, 
integrated with 6 subcutaneous bars on lower portion of trunk; upper portion of 
postanal trunk with 5 subcutaneous bars, third upper bar contiguous with third 
and fourth lower bars, last upper bar forming small, circular, entirely subcutaneous 
central caudal peduncle spot; upper, and to lesser extent, lower subcutaneous bars 
may be obscure. Four subcutaneous bars ventrolaterally on belly, not joined across 
ventral midline, bars narrow except in gravid females where broader in upper 
portion; anteriormost belly bar just posterior to pectoral fin base and lower portion 
of fourth bar terminating at anus; first 2 belly bars merging dorsolaterally, merged 
bar followed posteriorly by another dorsolateral subcutaneous bar aligned with 
the third lower belly bar; fourth belly bar lacking dorsal extension; 2 additional 
subcutaneous bars in nape region. Trunk mostly pale, faintly pigmented with fine 
scattered chromatophores; scale pockets usually unpigmented, rarely with fine 
peppery pigmentation along margins. 


VOLUME 96, NUMBER 4 791 


Dorsal fins usually uniformly pale to dark dusky although occasionally spinous 
dorsal fin pale with dusky bands; second dorsal fin irregularly dusky, sometimes 
with pale spots; anal fin uniformly moderate to dark dusky, often slightly darker 
than dorsal fins; caudal fin irregularly pigmented, usually pale or light dusky, 
sometimes with small dark spots on rays; pectoral and pelvic fins very slightly 
pigmented if at all. 

Geographic distribution. — Known from several localities in southern Japan and 
the Ryukyu Islands. 

Etymology.—The specific name japonica is based on the occurrence of this 
species in Japanese waters. 

Remarks.—Eviota japonica is a member of Group II (Lachner and Karnella 
1980:114) and is most closely related to E. queenslandica in that they share similar 
meristic characters, cephalic sensory pore patterns, and prominent patterns of 
dark spots on the head and pectoral base. They differ in the following details of 
the color pattern: the dark occipital spots of E. japonica tend to be irregular and 
angular in shape, and the spots on remainder of head somewhat elongate, but in 
E. queenslandica all spots are usually roundish; occipital spots of E. japonica are 
very dark, usually solid brown, at least in lower portions of spots, and usually 
darker than spots on cheek, opercle and sometimes pectoral base, whereas all 
spots in E. queenslandica are about equal in intensity to each other, the occipital 
as well as other spots composed of clusters of large, dark chromatophores rather 
than solid brown pigmentation; the branchiostegal membranes, lower margins of 
opercles, and the chin in E. japonica are usually heavily pigmented with elongate 
or angular dark marks, sometimes nearly as dark as occipital spots, but in E. 
queenslandica these areas have faint to moderate spots or scattered pigmentation, 
similar to the remainder of the head but less intensely pigmented; the trunk is 
mostly pale in E. japonica, at most with faint peppery chromatophores marking 
scale pockets, whereas the trunk in E. queenslandica is brownish due to heavier 
scale and scale pocket pigmentation; E. japonica has four subcutaneous bars in 
belly region, five subcutaneous bars above and six below on trunk posterior to 
anal fin origin, and with six postanal midline spots, and E. queenslandica has 
three bars or scattered chromatophores on belly region, four subcutaneous bars 
above and five below on posterior trunk, and with 5 postanal midline spots; the 
subcutaneous caudal peduncle spot in E. japonica is small, very faint and entirely 
subcutaneous whereas spot in FE. gueenslandica is small to moderate in size, 
somewhat more pronounced, and mostly subcutaneous, but often with a slight 
surface pigmentation as well. 

The above color characters distinguishing these two species show no intergra- 
dation in the Ryukyu Islands where the two species are sympatric. We now report 
E. queenslandica for the first time from the Ryukyu Islands (YCM-P2523, 2 
specimens, YCM-P4166, 2, YCM-P4069, 8, all from Ishigakijima Island; YCM- 
P2926, 2, from Taketomijima Island). 


Eviota latifasciata, new species 
Fig. 5 


Material examined.—88 specimens from 4 localities in the western Pacific and 
Indian Oceans, total size range 7.7—14.7, gravid females 10.7—13.2. 

Holotype: AMS I.18051-001, (12.4), female; Gilbert Islands, Abaiang Atoll, off 
Bolton Point, 7.6-10.7 m, 10 Nov 1973, D. F. Hoese. 


792 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 5. Eviota latifasciata, Paratype, 10.9 mm SL, juvenile, Abaiang Atoll, USNM 260079. 


Paratypes: GILBERT ISLANDS, ABAIANG ATOLL: AMS I.18051-073, 17 
(9.4-14.0); BPBM 28959, 2 (11.4, 12.9); AMNH 55062, 2 (11.2, 12.3); same data 
as holotype. USNM 260079, 4 (10.3-13.0); Lagoon side of Teirin Is., D. F. Hoese. 
AMS 1I.18043-001, 15 (8.4—-13.8); CAS 52737, 2 (11.6, 12.4); ANSP 151949, 2 
(12.4, 13.4); Lagoon off Teirin Is., 7.6 m, D. F. Hoese. CAROLINE ISLANDS: 
USNM 225034, 2 (11.7, 12.8); Ponape, 7°35'’N, 158°11'50”E, 0-15.2 m, V. G. 
Springer 80-8. USNM 225033, 6 (11.1—12.6); Ponape, 7°01'59’N, 158°14’02’E, 
O-4.6 m, V. G. Springer 80-11. USMN 225032, 4 (9.5-13.0); Ponape, 7°01'N, 
158°19’E, 0-18.3 m, V. G. Springer 80-22. USNM 225037, 15 (7.7-12.0); Sen- 
yavin Is., Ant Atoll, 6°47'N, 157°54'24”E, 0-24.4 m, V. G. Springer 80-17. 

Non-type material: 8 specimens from Christmas Island in the Indian Ocean 
(all WAM material); 10 specimens from Kapingamarangi Atoll, Caroline Islands, 
all in poor condition and faded (all CAS material). 

Diagnosis.—Cephalic sensory pore system lacking IT pore; pectoral fin with 
some branched rays; dorsal/anal fin ray formula typically 8/8; no filamentous 
spines in first dorsal fin; pelvic fin typically I, 4 plus rudiment and with reduced 
membrane between rays; branches on fourth pelvic fin ray 7-11; body generally 
pale with small dark occipital spot and 4 broad dusky subcutaneous postanal 
trunk bars; postanal midline spots inconspicuous; large dark midcaudal peduncle 
spot present. Diminutive species, all specimens less than 15 mm SL. 

Description.— Dorsal fin VI-I, 7(1), VI-I, 8(29), VI-I, 9(2); anal fin I, 8(30), I, 
9(2); pectoral fin rays 15(2), 16(/4), 17(9), 18(7); pectoral fin rays 10-18 may be 
branched, 11-15 usually branched; pelvic fin I, 4 plus rudiment (20), I, 4 1/10(3); 
branches on fourth pelvic fin ray 7-11, average 9.3; segments between consecutive 
branches of fourth pelvic fin ray 0-2, average 1.0; pelvic fin membrane reduced; 
branched caudal fin rays 11(7), 12(4); segmented caudal fin rays 17(20); lateral 
scale rows 23(3), 24(4), 25(1); transverse scale rows 6(6), 7(2); breast scaleless; 
vertebrae 10(S) precaudal plus 16(38) caudal, total 26. 

Spinous dorsal fin not elongate. Pelvic fin usually extending beyond origin of 
anal fin. Cephalic sensory pore system pattern 2; cutaneous papillae system not 
completely discernible in this small species. Male genital papilla not fimbriate. 

Color in preserved specimens.—Pale body, salient pigmentation consisting of 
small, dark occipital spot and 4 broad, dark, subcutaneous postanal trunk bars. 
Head mostly pale, occipital spot located laterally on head above midopercle, of 


VOLUME 96, NUMBER 4 793 


varying size and intensity, never large and pronounced. Some scattered chro- 
matophores dorsally on head just behind eyes. Fleshy base of pectoral fin unpig- 
mented. Trunk with 7-10 small, weak, rectangular shaped saddles over dorsum, 
from about middle of spinous dorsal fin posteriorly to caudal fin, better developed 
posteriorly, obscure in some specimens; belly with 2 broad, dusky subcutaneous 
patches extending dorsolaterally on trunk, and sometimes a faint subcutaneous 
bar dorsolaterally on nape; postanal portion of trunk with 4 broad, dusky to dark, 
subcutaneous bars, fourth either more intensely pigmented throughout or darker 
in midportion, appearing as large, dark, midcaudal peduncle spot overlying less 
dense subcutaneous bar. Postanal ventral midline spots inconspicuous. Spinous 
dorsal fin mostly pale, usually with a narrow horizontal dusky stripe basally and 
dusky distal margin; fin sometimes with scattered dark chromatophores through- 
out, never intensely dark; second dorsal fin similar to spinous dorsal fin, basal 
pigmentation more pronounced posteriorly. Anal fin pale to dark brown, usually 
darker than dorsal fin. Caudal fin pale to weakly pigmented, darker on lower half, 
sometimes rays with small dark spots. Pectoral and pelvic fins pale. 

In specimens from Ponape and Ant Atoll the subcutaneous trunk bars are mostly 
obscure and in some individuals the lower portion of the third postanal bar is 
partially divided resulting in five bars on the lower trunk. Specimens from Ka- 
pingamarangi Atoll are very faded, showing only the occipital and caudal peduncle 
spots. 

Geographic distribution.— This species is known from Christmas Island in the 
Indian Ocean, Kapingamarangi Atoll, Ant Atoll and Ponape, Caroline Islands, 
and from Abaiang Atoll, Gilbert Islands. 

Etymology.—The specific name is formed from the Latin words /atus, meaning 
broad or wide, and fasciatus, meaning banded, in reference to the broad subcu- 
taneous bars on the trunk. 

Remarks.—This species 1s a member of Group II (Lachner and Karnella 1980: 
114) and can be distinguished from other members of this group by the combi- 
nation of the following characters: dorsal/anal fin ray formula 8/8, no elongation 
of the spinous dorsal fin, a nonfimbriate male genital papilla, typically four post- 
anal subcutaneous trunk bars, the presence of an occipital spot, and an unpig- 
mented fleshy base of the pectoral fin. Eviota latifasciata shares several of the 
above characters with E. bipunctata and E. indica but differs from these in its 
coloration and reduced number of subcutaneous trunk bars. 

Some variation in the pectoral fin ray count of Eviota latifasciata with locality 
was observed; Christmas Island 17(2), 18(4), Kapingamarangi Atoll 17(4), 18(3), 
Ponape 15(2), 16(5) and Abaiang Atoll 16(9), 17(3). 


Eviota punctulata, new species 
Figs. 6—7 


Material examined.—255 specimens from 9 areas in Indonesia, Australia and 
western Oceania; total size range 7.3—23.1; gravid females 11.8—15.7. 

Holotype: USNM 224550, (20.7), male; Fiji Is., Great Astrolabe Reef, reef 
north of Vuro Is., 8 May 1965, R. Bolin and party, Te Vega Exp., Cr. 7, sta 278. 

Paratypes: FIJI ISLANDS: USNM 224543, 18 (18.0-23.1); same data as ho- 
lotype. USNM 224542, 3 (15.5-19.9); same locality as holotype, Te Vega Exp., 


794 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 6. Eviota punctulata, Paratype, 17.0 mm SL, male, Papua New Guinea, USNM 224548. 


Cr. 7, sta 279. PAPUA NEW GUINEA: USNM 224548, 18 (11.2-18.2); Ninigo 
Is., O0-9.1 m, V. G. Springer 78-3. USNM 224545, 1 (13.3); Hermit Is., O-O0.9 m, 
V. G. Springer 78-9. USNM 224547, 7 (11.6-17.8); Hermit Is., 0-7.6 m, V. G. 
Springer 78-18. USNM 224549, 17 (10.8-16.1); Hermit Is., 0-12 m, V. G. Spring- 
er 78-19. CAS 47909, 5 (13.0-17.0); AMS I.22213-001, 5 (11.2-16.9); above two 
lots with same data as USNM 224549. USNM 224544, 1 (17.8); New Ireland, 
Nusa Is., 0-10 m, T. Roberts. GREAT BARRIER REEF, AUSTRALIA: Lizard 
Is.: AMS I1.18740-107, 11 (14.8-18.0); Yonge Reef, D. F. Hoese, LZ 75-17. AMS 
1.18739-102, 3 (12.5-15.8); Palfrey Is., 3-10 m, D. F. Hoese, LZ 75-49. AMS 
1.20762-045, 1 (17.1); S. of Rocky Point, 1.5—4.6 m, D. F. Hoese 75-38. AMS 
1.21539-082, 1 (12.6); between Bird and South Is., 1.5—7.6 m, D. F. Hoese 75- 
113. AMS I.21343-008, 4 (10.1—13.8); Palfrey Is., 3 m, D. F. Hoese 75-5. USNM 
224546, 4 (11.4—20.4); Palfrey Is., 2-6 m, G. Anderson and B. Russell, FT 74-2. 
Endeavour Reef, collected by C. L. Smith and J. Tyler: ANSP 146495, 1 (16.4); 
13.7-18.3 m, TS,A-15. ANSP 146494, 4 (12.9-18.6); 7.6-18.3 m, TS,A-3. CAS 
47911, 3 (11.0-18.9); 13.7-15.2 m, TS,A-16. ANSP 146492, 4 (13.2-15.3); 13.7- 
19.8 m, TS,A-11. AMNH 42919, 7 (11.1-22.8); 0-13.7 m, S69-16. Little Hope 
Island, collected by C. L. Smith and J. Tyler: AMNH 42918, 1 (15.1); S69-28. 
AMNH 42920, 1 (18.8), 0-4.6 m, S69-30. PHILIPPINE ISLANDS: collected by 
the Smithsonian BFAR, and Silliman University teams, and Alcala: CAS 47910, 
3 (13.9-16.0); Palawan Prov., Bararin Is., O0-13.7 m, SP 78-20. USNM 224541, 
4 (15.3-18.3); Palawan Prov., Cuyo Is., Tagauayan Is., 0—-2.4 m, SP 78-25. AMS 
1.22209-001, 2 (15.1, 18.3); Palawan Prov., Cuyo Is., 0.6-—1.2 m, SP 78-17. 

Non-type material: Numerous specimens from the following general localities: 
INDONESIA; GREAT BARRIER REEF, AUSTRALIA; OCEANIA: Palau Is- 
lands; Kapingamarangi Atoll; Ponape Islands. 

Diagnosis.—Cephalic sensory pore system lacking IT pore; pectoral fin with 
some branched rays; dorsal/anal fin ray formula typically 9/8; elongation of spines 
in first dorsal fin rare; pelvic fin I, 4 1/10—2/10 with reduced membrane between 
rays; branches on fourth pelvic fin ray. 8-18; trunk with 6 dark postanal midline 
spots; head with dark, somewhat teardrop-shaped, postocular spot; midportion 
of fleshy base of pectoral fin with oblique streak of dusky pigmentation; scale 
pocket pigmentation on trunk well developed, more so posteriorly; dorsal and 
caudal fins with numerous prominent small dark spots. 

Description. — Dorsal fin VI-I, 8(1), VI-I, 9(23), VI-I, 10(1), VII-I, 9(1); anal fin 


Fig. 7. Eviota punctulata, Holotype, 20.7 mm SL, male, Fiji Islands, USNM 224550. 


I, 7(2), I, 8(24); pectoral fin 15(5), 16(16), 17(4), 18(/); pectoral fin rays 8-16 may 
be branched, 10-15 usually branched; pelvic fin I, 4 1/10(9), I, 4 2/10(/5); branches 
on fourth pelvic fin ray 8-18, average 13.7; segments between consecutive branch- 
es of fourth pelvic fin ray O—4, average 1.2; pelvic fin membrane reduced; branched 
caudal fin rays 12(14), 13(6), 14(2); segmented caudal fin rays 16(2), 17(24); lateral 
scale rows 23(7), 24(8), 25(5), 26(1); transverse scale rows 6(6), 7(/3); breast 
scaleless; vertebrae 10(/0) precaudal plus 16(/0) caudal, total 26. 

Spinous dorsal fin elongation rare and not well developed, only observed in 
males; first spine may be filamentous, and extending, when depressed, to base of 
fifth dorsal fin ray. Pelvic fin usually extending beyond origin of anal fin. Cephalic 
sensory pore system pattern 2; cutaneous papillae system pattern B. Male genital 
papilla not fimbriate. 

Color in preserved specimens.—Salient coloration of Eviota punctulata consist- 
ing of dark, boldly marked scale pockets, small distinct dark spots on dorsal and 
caudal fins, and oblong to teardrop-shaped mark dorsolaterally on head behind 
upper portion of eye. 

Cheek and preopercle with about 4—5 irregularly shaped patches of chromato- 
phores, one of which always more or less vertically elongate and occurring just 
posterior to rictus; opercle mostly pale, faint brownish patch on upper portion; 
snout more or less dusky; usually faint bar from eye to upper lip at about seven 
o’clock; chin and branchiostegal area with faint patches of chromatophores or 
dusky. Dark, more or less teardrop-shaped spot dorsolaterally on head behind 
upper portion of eye, spot darkest ventrally; spot extending obliquely toward 
dorsal midline, nearly meeting contralateral mark. Two smaller dark marks an- 
terior to these, on dorsal portion of head immediately behind eyes. Three or 4 
irregular transverse bars on nape, bars meeting at midline posterior to teardrop 
mark; bars less intense than teardrop spot and sometimes subdivided into irregular 
marks. 

Fleshy base of pectoral fin usually with oblique dusky mark through midlateral 
portion; pale areas above and below mark may be encircled by chromatophores; 
pectoral fin base frequently uniformly pigmented or with fine chromatophores 
above and below larger chromatophores in midlateral portion. Trunk character- 
istically pigmented with dark, vertically elongate, rather wide marks on scale 
pockets, these smaller on belly and most intensely pigmented posteriorly on trunk. 


796 PROCEEDINGS OF THE BIGLOGICAL SOCIETY OF WASHINGTON 


Belly with 3 dark, narrowly separated subcutaneous patches aligned with fainter 
subcutaneous patches located dorsolaterally on trunk; another dorsolateral sub- 
cutaneous patch anteriorly, on nape. Six postanal ventral midline spots, spots 
poorly developed and sometimes obscure, occasionally followed by weaker, small- 
er seventh spot near procurrent caudal rays. Six postanal spots associated with 
short, dark, subcutaneous bars usually faint or obscure. Seventh spot, when pres- 
ent, not confluent with bar although sometimes associated with very small sub- 
cutaneous marking. Upper subcutaneous trunk bars indistinct and rarely percep- 
tible, reduced in development. 

First dorsal fin variable, usually dark dusky brown on membrane, sometimes 
darker distally, base coloration of spines pale, spines with small dense black to 
brown spots: 4—5 spots occurring on first spine, spots fewer and sometimes fainter 
on subsequent spines, spots sometimes diffuse, indistinct or wanting; spots may 
be arranged in rows appearing as narrow oblique bars on fin. Membrane of second 
dorsal fin usually uniformly dusky brown but may be darker basally and distally, 
rays transparent with about 3—5 small discrete dark brown spots on each element. 
Caudal fin membrane dusky brown, rays mostly hyaline with about 6-8 small 
dark spots on each element, spots sometimes arranged to form irregular, vertically 
linear pattern. Anal fin uniformly dark brown, darker than second dorsal or caudal 
fins, equal to or slightly darker than first dorsal fin. Pectoral fin pale with very 
fine dark chromatophores bordering rays. Pelvic fin pale. 

Sexual dichromatism not pronounced. Females tending to be frayed and paler 
than males; most females with less pronounced spots on fins and scale pocket 
markings, although in some specimens marks equal in intensity to those on males. 

Geographic distribution.—This species occurs in the Java Sea and northward 
to the Philippine Islands, eastward to Ponape and Fiji Islands, and on the Great 
Barrier Reef, Australia. 

Etymology.—The specific name punctulata is derived from the Latin word 
punctum, and is used in reference to the small dark spots on the fins. 

Remarks.—Eviota punctulata is a member of Group II (Lachner and Karnella 
1980:114) and is easily distinguished from other members of the group in having 
a unique color pattern, extensive branching on the fourth pelvic fin ray and little 
or no elongation of the spinous dorsal fin. 


Eviota cometa, new species 
Figs. 8-9 


Material examined.—71 specimens from 4 general areas; total size range 10.3- 
18.5; gravid females 11.6—-14.8. 

Holotype: USNM 235817, 1 (15.7); Fiji Islands, Totoya Is., 18°58’57’S, 
179°52'12"W, 30 m, 27 Apr 1983, V. G. Springer, VGS 82-8A. 

Paratypes: FIJI ISLANDS: AMS 1.24027-001, 1 (14.5); Naviti Is., 17°06’S, 
177°13’E, 16.8-30 m, V. G. Springer 82-32. USNM 235832, 8 (10.3-13.7); ANSP 
151996; 2 (11.7, 13:3): CAS 52831, 2 3.2, 14:9): 18°42'S, 1789297 B= 0-390 etenae 
VGS 82-25. USNM 235863, 1 (13.0); Kandavu, 19°04’S, 178°02’E, 0-13.7 m, 
VGS 82-22. USNM 260328, | (16.4); N of Vuro Is., 18°52’S, 178°30.5’E, R. Bolin. 

Non-type material: Numerous specimens from four regions in the Pacific Ocean, 
the Ponape, Gilbert, Phoenix and Line Islands. 


VOLUME 96, NUMBER 4 797 


Fig. 8. Eviota cometa,/11.6 mm SL, male, Northern Escape Reef, Great Barrier Reef, Australia, 
ANSP 152010. 


Diagnosis.—Cephalic sensory pore system lacking IT pore; pectoral fin rays 
unbranched; dorsal/anal fin ray formula usually 8/7 or 9/8; spinous dorsal fin 
may contain filamentous spines, uncommon in females; pelvic fin typically I, 4 
1/10 with reduced to well developed membrane between rays; branches on fourth 
pelvic fin ray 4-9; body pale with prominent dark double mark at base of caudal 
fin and dark streak from mark along lower portion of caudal fin; subcutaneous 
bars and postanal midline spots absent in almost all specimens. 

Description.— Dorsal fin VI-I, 7(1), VI-I, 8(22), VI-I, 9(2/), VI-I, 10(1); anal 
fin I, 6(2), I, 7(21), I, 8(22); pectoral fin rays 13(1), 14(2), 15(7), 16(/2), 17(4); 
pelvic fin I, 4 plus rudiment (3), I, 4 1/10(17), I, 4 2/10(2); branches on fourth 
pelvic fin ray 4—9, average 6.7; segments between consecutive branches of fourth 
pelvic fin ray 1-5, average 1.7; pelvic fin membrane reduced to well developed, 
most often reduced; branched caudal fin rays 11(/4), 12(3), 13(1); segmented 
caudal fin rays 16(2), 17(24); lateral scale rows 21(1), 22(8), 23(2); transverse scale 
rows 5(7), 6(3), 7(1); breast scaleless; vertebrae 10(14) precaudal plus 15(14) 
caudal, total 25. 

First 3 spines of spinous dorsal fin in males may be filamentous, first longest, 
maximum extension to midcaudal peduncle; females rarely with slight elongation 
of first spine. Pelvic fin almost always extending beyond origin of anal fin. Cephalic 
sensory pore system pattern 2; cutaneous papillae pattern not well developed in 
this species. Male genital papilla not fimbriate. 

Color in preserved specimens. —Head and trunk mostly pale with dusky anterior 
nostrils, few scattered chromatophores above and behind eye, large scattered 
subcutaneous chromatophores on belly. Prominent dark, basicaudal mark con- 
sisting of 2 portions: roundish spot, about size of pupil, midlaterally at end of 
caudal peduncle, touching or nearly so, a dark, elongate, vertical mark at base of 
caudal rays; vertical mark extending from slightly above midline nearly to ventral 
margin of fin. Dark, dusky streak extending posteriorly from lower half of dark 
vertical mark to distal margin of fin; remainder of caudal fin mostly pale. Spinous 
dorsal fin with dusky band basally, otherwise pale. Second dorsal fin with dusky 
band basally and distally, or in large males, dusky throughout. Anal fin dusky on 
lower half or dusky throughout, slightly darker than other fins. Pectoral and pelvic 
fins pale, except in large males where they are faintly dusky. 

Postanal midline of trunk pale or with many tiny subcutaneous spots, usually 


798 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 9. Eviota cometa, 15.6 mm SL, male, Palmyra, CAS 52852. 


not forming distinct large spots; few specimens from Line Islands having 5 distinct 
postanal midline spots. Belly with large scattered chromatophores, lacks distinct 
bars or pigment patches. 

In specimens from Fiji, scale pockets along ventroposterior portion of trunk 
weakly pigmented, pigmentation consisting of single row of chromatophores bor- 
dering each scale pocket; this pigmentation may also occur more dorsally on 
trunk, but less well developed. 

Large male specimens having darker fins and somewhat enlarged, more diffuse, 
basicaudal mark. Males may also have broad faint dusky band through lower two- 
thirds of head and trunk; band consisting of fine, widely scattered, dark chro- 
matophores. Males from Abaiang Atoll having enlarged basicaudal spot, anterior 
portion of which deeper than in other populations and nearly merging with pos- 
terior portion, latter also broader than typical. Females from Abaiang having 
more typically shaped mark. 

Color in life.—The following description is from a color slide of a 16.1 mm SL 
male specimen, collected at Fanning Island, 20 Sep 1978, by P. S. Lobel (Northern 
Territory Museum, Australia). Body pale with reddish-brown pigmentation through 
most of lower portion of trunk, pigmentation beginning at eye, extending across 
upper cheek and opercle, over belly region where intensity greatest, terminating 
at end of caudal peduncle where coloration weakest; eye and anterior nostrils with 
some reddish pigmentation. Lower half of head with some scattered, fine, dark 
chromatophores. White spots within reddish-brown coloration on body: 3 or 4 
spots laterally on head, 5 or 6 on belly, 6 on postanal ventral midline of trunk, 
those on belly most distinct, on posterior trunk most diffuse. Six elongate white 
spots bordering upper limit of reddish-brown coloration just above midlateral 
body septum. Large black basicaudal mark at hypural joint, anterior roundish 
part merging somewhat with posterior bar-like portion, margins of both sections 
diffuse. Weak dusky streak extending from lower portion of basicaudal spot hor- 
izontally through lower portion of caudal fin to distal margin of fin. Second dorsal 
and anal fins with slight dusky pigmentation posteriorly, otherwise pale. All other 
fins pale, lacking prominent pigmentation. 


VOLUME 96, NUMBER 4 799 


Geographic distribution.— This species occurs in the Great Barrier Reef, Aus- 
tralia, and Oceania at Fiji, Ponape and the Gilbert, Phoenix and Line Islands. 

Etymology.—The specific name cometa is Latin for comet and is used in ref- 
erence to the basicaudal spot and the trailing dark streak. 

Remarks.—Eviota cometa is a member of Group III (Lachner and Karnella 
1980:114). Of the species in this group it most closely resembles E. zebrina. Both 
species share a variable dorsal/anal fin ray formula of 8/7 or 9/8, have a pelvic 
fin membrane that ranges in development from reduced to well developed, and 
have similar prominent basicaudal spots. Eviota cometa differs from E. zebrina 
in lacking well developed body pigmentation, other than the basicaudal mark, 
and lacking vertical bars on the caudal fin. Eviota cometa has a dark horizontal 
streak on the lower portion of the caudal fin not found in specimens of E. zebrina 
from Oceania and Australia. . 

Eviota cometa and E. zebrina are sympatric at Fiji and the Great Barrier Reef. 
At Fiji most specimens of both species have a dorsal/anal fin ray formula of 9/8 
but on the Great Barrier Reef E. zebrina has a formula of 9/8 and E. cometa 
8/7. 

No pattern emerges when the dorsal/anal fin ray formula of Eviota cometa is 
plotted against geographic locality. The formula varies within and between lo- 
calities. 


Eviota sigillata, new species 
Fig. 10 


Material examined.—490 specimens from several areas extending from the 
Indian Ocean to Oceania; total size range 9.0—21.0; gravid females 12.4—15.0. 

Holotype: USNM 223836, (18.3), male; Indian Ocean, St. Brandon Shoals, 
south of Raphael Is., O0-3.7 m, 8 Apr 1976. V. G. Springer, VGS 76-12. 

Paratypes: INDIAN OCEAN, ST. BRANDON SHOALS: USNM 223834, 31 
(10.3-17.7); CAS 47938, 5 (14.0-18.1); AMS I.22205-001, 5 (13.6-18.0); ANSP 
146761, 5 (13.1-16.8); BPBM 26538, 4 (14.1-18.6); WAM P.27053-001, 3 (12.0- 
17.7); ROM 36923, 3 (14.8-17.7). Preceding seven lots from same collection as 
holotype. USNM 223830, 1 (14.6); Albatross Is., 0-18.3 m, VGS 76-22. USNM 
223827, 4 (15.6-19.6); N of Frigate Is., 15.2-21.3 m, VGS 76-5. USNM 223831, 
29 (12.3-20.3); S of Raphael Is., 0-9.1 m, VGS 76-20. USNM 223835, 16 (9.6- 
19.0); southern part of Shoals, 15.2 m, VGS 76-18. USNM 223828, 25 (10.1- 
19.7); E of Raphael Is., 7.6-9.1 m, VGS 76-8. USNM 223833, | (14.4); E of 
Raphael Is., 0.2-1.1 ra, VGS 76-7. USNM 223829, 12 (10.9-18.1); 0-2 m, VGS 
76-1. USNM 223832, 7 (16.6—21.0); NE of Siren Is., 16.8-—21.3 m, VGS 76-19. 
RUSI 1888, 2 (13.6, 16.1); W of Tortue Is., T. H. Fraser, SA-35. 

Non-type material: Numerous specimens from the following general areas: IN- 
DIAN OCEAN: Seychelles Islands, Chagos Archipelago, Sri Lanka; INDONESIA; 
GREAT BARRIER REEF, AUSTRALIA; OCEANIA: Yap Island, Kapinga- 
marangi Atoll, Ponape Islands. 

Diagnosis.—Cephalic sensory pore system lacking IT pore; pectoral fin rays 
unbranched; dorsal/anal fin ray formula typically 9/8 with some geographic vari- 
ation to 8/7; spinous dorsal fin may contain filamentous spines in both sexes; 


800 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


b 


Fig. 10. E£viota sigillata. a, 13.0 mm SL, female, Sri Lanka, USNM 223841; b, 17.4 mm SL, male, 
Seychelles Islands, ANSP 146506. 


pelvic fin I, 4 1/10—2/10 and with well developed membrane between rays; branch- 
es on fourth pelvic fin ray 3-7; trunk with 7 dark postanal midline spots; coloration 
variable with age and sex: females and small males with chromatophores some- 
what clustered laterally on head, 2 dark spots basally on upper and lower portions 
of caudal fin and additional weaker spots on fin membrane, and with dusky 
pigmentation basally in dorsal fins; large males, frequently more robust in body 
shape, having uniformly distributed chromatophores on head, more diffuse spot- 
ting on caudal fin, lower basicaudal spot more prominent, and dorsal fins with 
more extensive dusky pigmentation. 

Description.— Dorsal fin VI-I, 8(7), VI-I, 9(23), VI-O 10(1); anal fin I, 7(7), I, 
8(24); pectoral fin rays 14(1), 16(9), 17(70), 18(10), 19(1); pelvic fin I, 4 1/10(/5), 
I, 4 2/10(16); branches on fourth pelvic fin ray 3—7, average 5.3; segments between 
consecutive branches of fourth pelvic fin ray 1-7, average 3.4; pelvic fin membrane 
well developed; branched caudal fin rays 11(2/); segmented caudal fin rays 17(3 1); 
lateral scale rows 21(6), 22(4), 23(3), 24(1); transverse scale rows 5(1), 6(5); breast 
almost always scaleless, single embedded scale observed in one specimen; pre- 
caudal vertebrae 10(20), caudal vertebrae 15(/9), 16(1), total 25(/9), 26(1). 

First 4 dorsal fin spines of males and first 3 spines of females may be filamentous, 
spines longer in males, when depressed, longest spine extending to procurrent 
caudal fin rays. Pelvic fin always extending beyond origin of anal fin. Cephalic 
sensory pore system pattern 2; cutaneous papillae system pattern B. Male genital 
papilla not fimbriate. 

Some mature males may have deeper bodies than females and immature males. 
Head depth, at posterior margin of opercle, expressed as thousandths of the 
standard length, 213-234 (7 spec. 13.3-20.1 mm SL) for deep bodied or stout 


VOLUME 96, NUMBER 4 801 


males; 201-215 (6 spec., 16.3-19.7) for nonstout males; and 209-219 (6 spec. 
13.4-16.0) for females. 

Color in preserved specimens.— There is markedly pronounced sexual dimor- 
phism in coloration and body depth. In mature females and nearly all small 
specimens of both sexes (less than approximately 13-17 mm SL), dark color 
laterally on head irregular and somewhat clustered, most concentrated on anterior 
portion of opercle, cheek pale or with few chromatophores, and snout and nostrils 
very dark, prominent against pale upper jaw and chin. Head dorsally and nape 
with few small dark spots, usually along midline, followed by series of 11 to 13 
small dark spots on midline of trunk from origin of spinous dorsal fin to end of 
caudal peduncle, latter series faint or obscure in many specimens. Fleshy base of 
pectoral fin variably pigmented, often with small oblique patch of chromatophores 
in central portion, or with patches on upper or lower portions, base entirely pale 
in many specimens. Scale pockets weakly pigmented if at all, development of 
pigmentation mostly restricted to upper anterior portion of trunk, remainder of 
trunk predominantly pale. Seven small, dark, postanal midline spots integrated 
with 7 narrow dark vertical subcutaneous bars extending to slightly below mid- 
trunk, pale interspaces usually wider than dark bars; subcutaneous bars on upper 
trunk obscure. Two broad, oblique, dark subcutaneous patches on belly. First and 
second dorsal fins with dusky pigmentation basally, outer portions pale. Anal fin 
light dusky or pale. Caudal fin with 2 dark spots, on upper and lower basal portions, 
lower spot larger and more prominent; 6 to 8 smaller spots scattered over rest of 
fin, often weakly developed, few specimens with spots arranged in 2 or 3 wavy 
vertical bands; lower portion of caudal fin dusky from base to distal margin. 
Pectoral and pelvic fins unpigmented. 

Small and some moderate sized males have pigment patterns nearly identical 
to females, although somewhat darker. As males mature they develop a color 
pattern that differs from females and all juveniles in the following ways: pigmen- 
tation laterally on head, including snout, cheek, and opercle, more or less uni- 
formly distributed rather than clustered as on anterior portion of opercle of females 
and juveniles, and consisting of small evenly spaced dark brown chromatophores; 
upper caudal fin spot less intense and lower spot larger and more diffuse; dark 
basal pigmentation of spinous and second dorsal fins is broader and darker, 
extending to distal margins of fins in some specimens; postanal midline spots 
larger and more diffuse, sometimes seemingly merging with one another; dark 
subcutaneous bars on posterior lower trunk usually obscure. 

Males also undergo dimorphism in head and body depth with maturity, evident 
as deepening of anterior part of body (Fig. 10b). This always correlated with 
mature male color pattern found in specimens that range in size from approxi- 
mately 13 to 21 mm SL. Males with deep bodies much less common and usually 
among largest specimens in a given collection. Size at which dimorphism occurs 
varies among collections and with localities. 

Our description of the color in preservation is based mostly on specimens from 
the St. Brandon Shoals. Almost all other specimens were faded to some degree 
and do not exhibit the full color pattern as seen on most St. Brandon Shoals 
specimens. The holotype is a male representing a transitional stage of color pattern 
between that of the juvenile and large adult male. 

Color in life. —The following color description is based on a color transparency 


802 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


of a 14.4 mm SL female captured in the Chagos Archipelago on March 5, 1979 
by R. Winterbottom and A. R. Emery (ROM 36877). Snout, excluding upper lip, 
orange-red; iris golden orange-red with some white in upper portion, pupil black; 
head from eye to end of opercle reddish-orange, becoming light golden orange 
dorsally on head and nape; upper and lower jaws, chin, head below eye, and fleshy 
base of pectoral fin pale. The two dark oblique subcutaneous marks found on 
belly of preserved specimens golden-orange, separated by milky-white areas. Elev- 
en or 12 small orange spots on trunk along dorsal midline beginning at origin of 
spinous dorsal fin, preceded by 3 small orange spots dorsally on head and nape. 
Trunk dorsolaterally, especially anteriorly, light orangish; scale pockets, above 
midline of body, reddish-orange. Seven subcutaneous spots and bars along ventral 
midline reddish-orange, lower portions of bars most intense. Two reddish-orange, 
vertically aligned spots, basally on upper and lower portions of caudal fin, lower 
spot somewhat larger. Small faint orangish spots elsewhere on caudal fin and 
small, milky-white spots on rays with tiny whitish spots also on membrane of 
lower portion of fin. First and second dorsal fins with very small faint orangish 
spots, basal portions with tiny milky-white spots; tiny milky-white spots also on 
pelvic and anal fins. 

Geographic distribution.—Eviota sigillata is known from several localities in 
the Indian Ocean, a single locality in Indonesia, several localities in the northern 
portion of the Great Barrier Reef, Australia, and from Yap, Palau and Ponape 
Islands of Oceania. 

Etymology.—The specific name is derived from the Latin word sigi//atus, mean- 
ing adorned with little figures or marks, in reference to the seven dark, subcuta- 
neous ventral midline spots. 

Remarks.—This species is a member of group III (Lachner and Karnella 1980: 
114) and aspects of its color pattern may resemble those of E. storthynx and E. 
zebrina, also of this group. Eviota sigillata differs from E. storthynx in lacking a 
dark, postocular spot, and differs from E. zebrina in having dark spots on upper 
and lower portions of caudal fin base, rather than a single central dark spot. 

Geographic differentiation occurs in two meristic characters, the dorsal/anal fin 
ray formula and the number of pectoral fin rays. Specimens from the Great Barrier 
Reef and Indonesia have a dorsal/anal fin ray formula of 8/7(6 specimens), and 
have 14—17(6) pectoral fin rays, whereas specimens from the Indian Ocean and 
Oceania typically have a formula of 9/8 (22 of 24 specimens) and have 16—19(24) 
pectoral fin rays. 


Eviota sparsa, new species 
Figs. 11-13 


Material examined. —215 specimens from 5 general areas; total size range 7.3- 
21.3; gravid female 15)9- 

Holotype: USNM 227483, 1 (16.9), male; Samoa Islands, Tutuila Is., at Utulei 
village, 15.2 m, R. Wass. 

Paratypes: SAMOA ISLANDS: USNM 260327, 7 (8.4-18.6); CAS 52832, 2 
(14.9, 16.3); ANSP 151998, 2 (13.6, 16.9); same data as holotype. PHILIPPINE 
ISLANDS: Collected by Smithsonian Philippine expeditions in 1978 and 1979: 
USNM 227485, 1 (13.4); Palawan Prov., Cuyo Is., Cocoro Is., 0-21.3 m, SP 78- 


VOLUME 96, NUMBER 4 803 


Fig. 11. Eviota sparsa, Holotype, 16.9 mm SL, male, Samoa Islands, USNM 227483. 


27. AMS 1.23987-001, 1 (17.2); Siquijor Is., 0-10.7 m, SP 78-7. AMNH 55055, 
1 (16.7); Palawan Prov., Cuyo Is., Tagauayan Is., 0-13.7 m, SP 78-24. USNM 
227481, 4 (15.6-17.9); Siquijor Is., 0-30.5 m, LK 79-16. INDONESIA: USNM 
210070, 3 (14.2-16.6); Saparua off Kampungmahu, 13.7-16.8 m, VGS 73-12. 
USNM 227484, 3 (15.3-15.9); Banda Islands, VGS 74-10 or 74-11. 

Non-type material: Numerous specimens from the following areas: Indonesia; 
Palau Islands; Great Barrier Reef, Australia. 

Diagnosis.— Cephalic sensory pore system lacking IT and POP pores; pectoral 
fin with some branched rays; dorsal/anal fin ray formula typically 9/8; elongation 
of spines in first dorsal fin uncommon; pelvic fin typically I, 4 6/10-8/10 with 
well developed membrane between rays; branches on fourth pelvic fin ray 3-5; 5 
dark postanal midline spots; body generally dusky brown; specimens from In- 
donesia, Philippine and Palau Islands may have prominent marks laterally on 
head. 

Description.— Dorsal fin VI-I, 8(3), VI-I, 9(26), VI-I, 10(1); anal fin I, 7(2), I, 
8(27); pectoral fin rays 14(1), 15(2), 16(/3), 17(7), 18(6); pectoral fin rays 8-17 
may be branched, 12-15 usually branched; pelvic fin I, 4 6/10(S), I, 4 7/10(16), 
I, 4 8/10(5), I, 4 9/10(1); 1 specimen in 30 with divided fifth pelvic fin ray; 
branches on fourth pelvic fin ray 3—5, average 4.1; segments between consecutive 
branches of fourth pelvic fin ray 2-8, average 4.5; pelvic fin membrane well 
developed; branched caudal fin rays 11(1), 12(5), 13(12), 14(2), 15(2); segmented 
caudal fin rays 16(1), 17(28), 18(1); lateral scale rows 23(4), 24(22), 25(3); trans- 
verse scale rows 6(10), 7(17); breast scaleless; vertebrae 10(//) precaudal plus 
16(//) caudal, total 26. 

Elongation of spinous dorsal fin uncommon, first 4 spines in males may be 
filamentous, longest spine extending to about middle of base of second dorsal fin; 
first 2 spines of spinous dorsal fin in females rarely slightly elongate, never fila- 
mentous. Pelvic fin usually reaching origin of anal fin or beyond. Cephalic sensory 
pore system pattern 5; cutaneous papillae system pattern B-1, as found in £. 
storthynx (Lachner and Karnella 1980:7). Male genital papilla not fimbriate. 

Color in preserved specimens. — Body more or less uniformly dusky brown, most 
specimens lacking accentuated color markings. Pigmentation of head laterally 
ranging from scattered dark brown chromatophores to variously intense clusters 


804 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 12. Eviota sparsa, 15.3 mm SL, male, Northern Escape Reef, Great Barrier Reef, Australia, 
ANSP 148483. 


of chromatophores, 2 most prominent clusters occur dorsolaterally behind eye 
and above preopercle. These patches more or less vertically elongate and rect- 
angular shaped, anterior cluster smaller. Patches sometimes integrated with weak- 
er transverse bars or spots on dorsal portion of head. These clusters usually well 
developed in specimens from Indonesia, Philippines and Palau Islands, moderate 
to weak in Samoan specimens, and weak or obscure in Australian specimens. 
Cheek pigmentation varying from scattered chromatophores to weak or moder- 
ately dark clusters of chromatophores; 2 to 5 clusters present, 2 of which may 
radiate from lower eye. Chin and snout with fine scattered chromatophores. Nape 
pigmentation ranging from uniformly scattered chromatophores to 2 or 3 weak, 
irregular transverse bars or irregular wavy clusters of chromatophores. 

Fleshy base of pectoral fin with scattered chromatophores or varyingly elongate 
aggregations of chromatophores on lower, or lower and upper portions of base, 
separated by pale or less intensely pigmented area. Sometimes a weak vertical 
dusky band through basal portion of pectoral fin; band may be integrated with 
one or both elongate pigment patches. Trunk with brownish crescent shaped marks 
on scale pockets, marks composed of 2 or 3 rows of chromatophores; scale pocket 
pigmentation forms a rather uniform diamond pattern over trunk. Five somewhat 


Fig. 13. Eviota sparsa, Paratype, 15.7 mm SL, female, Saparua, Indonesia, USNM 210070. 


VOLUME 96, NUMBER 4 805 


elongate, dark subcutaneous postanal midline spots integrated with 5 bars on 
posteroventral region of trunk, bars weak to moderately developed, sometimes 
obscure; posterodorsal portion of trunk with faint or obscure subcutaneous bars. 
Sixth dark ventral midline spot may occur at base of procurrent rays, often obscure. 
No prominent dark external or subcutaneous midcaudal peduncle spot. Belly with 
3 fairly wide subcutaneous bands, first 2 extending vertically entire depth of trunk, 
third occurring on lower trunk only, bands usually faint to obscure. 

Spinous dorsal fin most often uniform brownish but may have scattered dusky 
blotches or a dusky basal band. Membrane of anal and second dorsal fins uniformly 
light dusky, basal portions may be slightly darker, fin rays pale; some specimens 
with few pale circular spots on second dorsal fin; some specimens with anal fin 
slightly darker than dorsal fins. Caudal fin pale to light dusky, sometimes with 
several tiny weak dark spots on rays. Pectoral and pelvic fins pale. 

Color in life. —The following color notes were recorded by Richard Wass from 
an undetermined specimen of the type series, collected at 15.2 meters, Utulei 
village, Tutuila Is., Samoa: “Background coloration pale. Posterior halves of scales 
covered with tiny orange and yellow spots outlined in dusky resulting in brown 
effect at a distance. Dorsal and caudal rays with reddish orange spots. Anal rays 
red. Fin membranes dusky. Three darkly pigmented (internal) areas at base of 
anal and three on ventral portion of caudal peduncle. Orange spots on lower lip 
and chin. Dusky orange spots on cheek and nape.” 

Geographic distribution.—This species is known from Indonesia, the Philip- 
pines, Palau Islands, the Great Barrier Reef, and the Samoan Islands. 

Etymology.—The specific name sparsa is a Latin word meaning sprinkled or 
flecked, in reference to the chromatophores scattered over the body. 

Remarks.—Eviota sparsa is a member of Group VII. It can be distinguished 
from the other member of this group by the following characters: a long fifth 
pelvic fin ray and a uniform, unaccentuated coloration in most specimens. 


Acknowledgments 


We wish to extend our thanks to all those who participated in making specimens 
available to us on loan. In addition, we thank T. H. Fraser and R. Wass for 
descriptive accounts of liye coloration of specimens, and R. Winterbottom, A. 
Emery, P. Lobel, and Helen Larson for color slides that contributed to live color 
descriptions. The following individuals generously allowed us to make specimen 
exchanges in order to distribute paratypes among major museums: J. E. Bohlke, 
C. L. Smith, and D. F. Hoese. Finally, special thanks go to M. Hayashi (YCM), 
C. L. Smith (AMNH) and R. Wass (Office of Marine Resources, Government of 
American Samoa), for allowing us to retain specimens from their collections as 
holotypes at the USNM. 

The specimens illustrated in Fig. 1a and Fig. 10b were photographed by Kjell 
Sandved. The drawings shown in Figs. 2, 5, 7 and 9 were rendered by J. R. 
Schroeder, and those in Figs. 4, 12 and 13 by Paul Mazer. 


Literature Cited 


Karnella, S. J., and E. A. Lachner. 1981. Three new species of the Eviota epiphanes group having 
vertical trunk bars (Pisces:Gobiidae).— Proceedings of the Biological Society of Washington 
94(1):264—275. 


806 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Lachner, E. A., and S. J. Karnella. 1978. Fishes of the genus Eviota of the Red Sea with descriptions 
of three new species (Teleostei:Gobiidae). Smithsonian Contributions to Zoology 286:1—23. 
, and 1980. Fishes of the Indo-Pacific genus Eviota with descriptions of eight new 
species (Teleostei:Gobiidae).— Smithsonian Contributions to Zoology 315:1—127. 


Department of Vertebrate Zoology (Fishes), National Museum of Natural His- 
tory, Smithsonian Institution, Washington, D.C. 20560. 


PROC. BIOL. SOC. WASH. 
96(4), 1983, pp. 807-855 


REVISION OF THE GENUS LEPTOSOMATUM 
BASTIAN, 1865 (NEMATODA: LEPTOSOMATIDAE) 


Tom Bongers 


Abstract.—The available type-material of the species of Leptosomatum, has 
been studied and compared with the type-species of the genera Leptosomatides 
(L. euxinus Filipjev, 1918) and Syringonomus (S. typicus Hope and Murphy, 
1959). 

The character complex present in L. elongatum, the type-species of Leptoso- 
matum, sharply distinguishes the species of Leptosomatum from the genus Lep- 
tosomatides, hitherto regarded as being related to Leptosomatum. Females of 
Leptosomatides can be distinguished from those of Leptosomatum by the presence 
of a strongly muscularized vagina wall, here termed the vaginal ovejector. The 
main distinguishing character is the presence of a sexual dimorphism in the 
amphids of Leptosomatum, which is absent in Leptosomatides. 

The species of Leptosomatum can be grouped into three complexes: a) The 
monotypic complex L. kerguelense Platonova, 1958 (new synonyms: L. clavatum 
Platonova, 1958 partim, L. crassicutis Platonova, 1958, and L. arcticum sensu 
_ Mawson, 1958) characterized by the presence of a cephalic capsule in both sexes. 
b) The L. bacillatum-complex composed of L. bacillatum (Eberth, 1863) (new 
synonyms: L. elongatum Bastian, 1865, L. filipjevi Schuurmans Stekhoven, 1950, 
and L. tuapsense Sergeeva, 1973), L. sachalinense Platonova, 1978 (new synonym: 
L. diversum Platonova, 1978), L. acephalatum Chitwood, 1936 and probably 
L. clavatum Platonova, 1958 partim and L. sundaense n.sp. for L. sabangense 
sensu Micoletzky, 1930 nec Steiner, 1915. This complex is characterized by the 
presence of a cephalic capsule in juveniles and females, but not in males. c) The 
L. punctatum-complex with L. punctatum (Eberth, 1863) (new synonyms: L. 
longisetosum Schuurmans Stekhoven, 1943 and (?) Stenolaimus macrosoma Mar- 
ion, 1870), and L. keiense Micoletzky, 1930. In this complex the cephalic capsule 
is absent in juveniles and adults. 

More information is needed regarding the species L. abyssale Allgén, 1951; L. 
bathybium Allgén, 1954; L. behringicum Filipjev, 1916; L. breviceps Platonova, 
1967; L. groenlandicum Allgén, 1954; L. indicum Stewart, 1914; L. pedroense 
Allgén, 1947; L. sabangense Steiner, 1915; L. tetrophthalmum Ssaweljev, 1912 
and L. sundaense new name; pro L. sabangense sensu Micoletzky, 1930, they are 
considered species inquirendae. 

Leptosomatum ranjhai Timm, 1960, and L. micoletzkyi Inglis, 1970, do not 
belong to Leptosomatum and are, for the moment, considered species incertae 
sedis. 

L. caecum Ditlevsen, 1923 belongs to Pseudocella. 

L. arcticum Filipjev, 1916; L. elongatum sensu Platonova, 1967; L. gracile 
sensu Allgén, 1954; L. grebnickii Filipjev, 1916 and L. tetrophthalmum sensu 
Platonova, 1967 are transferred to Leptosomatides Filipjev, 1918. 


The genus Leptosomatum Bastian, 1865, which contains large-sized marine 
nematodes, was last revised by Filipjev (1918). Platonova (1976) published a key 


808 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


and reviewed the family Leptosomatidae. According to these authors the genus 
is characterized by the reduced cephalic capsule and the simple gubernaculum. 
On this basis, females of Leptosomatum cannot be distinguished from those of 
Leptosomatides Filipjev, 1918. The genus now contains 31 nominal species and 
identification of these has become impossible. 

While studying populations of a Leptosomatum species I observed some phe- 
nomena that appeared to be undescribed. Re-examination of type-specimens of 
most species revealed the presence of a character complex that clearly demarcates 
Leptosomatum from related genera. Some species that do not possess this complex 
had to be excluded from Leptosomatum, some were transferred to Leptosomatides 
and Pseudocella, and others must be regarded as species inquirenda. 

In a series of papers, starting with the present one, I will try to raise the clas- 
sification of the Leptosomatidae from the a-level and to establish a classification 
based on holapomorphy as proposed by Lorenzen (1981). The first step is the 
demarcation of the genera. Lorenzen did not succeed in basing the classification 
of the family Leptosomatidae on holapomorphy, partly because many species 
descriptions are incomplete and inadequate. 

Many species are synonymized, due to poor descriptions, sexual dimorphism, 
and post-adult growth. 

In this paper I give an historical account, call attention to artifacts, give sup- 
plementary descriptions, discuss the species, and provide a key. In another paper 
I shall give a phylogenetic approach. 


Historical Review 


The genus Leptosomatum was erected by Bastian (1865) who included eight 
species: L. punctatum (Eberth, 1863); L. gracile Bastian, 1865; L. bacillatum 
(Eberth, 1863); L. figuratum Bastian, 1865; L. coronatum (Eberth, 1863); L. 
longissimum (Eberth, 1863); L. subulatum (Eberth, 1863), and L. elongatum 
Bastian, 1865. The last mentioned was designated as “typical species.””! 

Marion (1870) transferred L. coronatum to Thoracostoma. Villot (1875) syn- 
onymized L. figuratum with L. coronatum, and described L. roscovianum, L. 
magnum and L. minutum. It was soon realized that these species did not conform 
to Bastian’s definition of the genus: de Man (1889) made L. magnum type-species 
of his new genus Cylicolaimus; in 1918 Filipjev transferred L. minutum to the 
same genus and in 1927 he removed L. roscovianum to Synonchus. 

Von Linstow, also, broadened the scope of Leptosomatum; he transferred Thor- 
acostoma schneideri (Biitschli, 1874) to it and described four new species: L. 
antarticum (1892), L. setosum (1896), L. papillatum (1903), and L. australe (1907); 
of these L. setosum was transferred to Thoracostoma by de Man (1904); L. 
antarticum and L. papillatum were removed to Deontostoma by Filipjev (1916), 
and L. australe was considered species inquirenda by Filipjev (1918). In 1893 de 
Man synonymized L. gracile with L. elongatum. 

The generic revision by Filipjev (1918) recognized only three of the above 
mentioned species, viz. L. bacillatum, L. punctatum and L. elongatum. In ad- 


! The indication “typical species” did not orginally have the nomenclatorial meaning it has today. 
According to Stiles and Hassal (1905), L. elongatum has to be considered as type by original desig- 
nation. 


VOLUME 96, NUMBER 4 809 


dition four other species were included: L. tetrophthalmum Ssaweljev, 1912; L. 
grebnickii, L. arcticum, and L. behringicum, the latter three previously described 
by Filipjev in 1916. In the same paper Filipjev erected the genus Leptosomatides. 
He was somewhat uncertain about the generic placement of L. grebnickii and L. 
arcticum because in some respects these species resembled the type-species of 
Leptosomatides, L. euxinus; but as males of these two species were still unknown, 
he left them in Leptosomatum. 

In 1936 Chitwood described L. elongatum subsp. acephalatum; in 1951 he 
reunited this form with the nominate form, but Timm (1953), in an anatomical 
and morphological study, raised it to specific rank. 

Allgén (1947, 1951, 1954, 1954a, and 1957) described five species, which are 
all doubtful, being described from single specimens. Moreover he confounded 
Leptosomatum and Leptosomatides; this will be discussed later. 

Platonova (1958, 1967, 1978) published on specimens of Leptosomatum iden- 
tified by Filipjev in the twenties, and described some new species. In her thesis 
(1976) she reviewed the genus and gave a key. 

Minor contributions to the taxonomy of Leptosomatum were made by Stewart 
(1914), Steiner (1915, 1916), Ditlevsen (1923), Kreis (1928), Micoletzky (1924, 
1930), Schuurmans Stekhoven (1943a, 1943b, 1950), Mawson (1958), Timm 
(1960), Inglis (1971), and Sergeeva (1973). 


Material and Methods 


The original material of the following species was studied: L. elongatum sensu 
de Man, 1893; L. arcticum, L. grebnickii and L. behringicum Filipjev, 1916; L. 
bacillatum (=L. filipjevi Schuurmans Stekhoven, 1950) and L. punctatum sensu 
Filipjev, 1918; L. coecum Ditlevsen, 1923; L. elongatum subsp. acephalatum 
Chitwood, 1936; L. sabangense sensu Allgén, 1942; L. acephalatum sensu Timm, 
1953; L. bathybium Allgén, 1954; L. groenlandicum Allgén, 1954; L. crassicutis, 
L. kerguelense and L. clavatum Platonova, 1958; L. ranjhai Timm, 1960; L. 
breviceps Platnonova, 1967; L. arcticum, L. elongatum and L. tetrophthalmum 
sensu Platonova, 1967; L. tuapsense Sergeeva, 1973; L. diversum and L. sach- 
.dlinense Platonova, 1978. 

Of related genera, type-specimens of Syringonomus typicus Hope and Murphy, 
1969 were studied, as well as the Leptosomatides collection of the Zoological 
Institute in Leningrad. 

Furthermore 80 specimens in the collection of the Smithsonian Institution, 
Washington, were made available; as well as some hundreds of specimens from 
the Dutch coast, deposited in the nematode collection of the Nematology De- 
partment, Landbouwhogeschool, Wageningen. These latter specimens had mainly 
been collected from the sponge Halichondria panicea (Pallas, 1766), in which they 
occur in great densities; from 100 ml of sponge more than 900 specimens were 
collected. The sponges were taken off stones in the lower littoral, and immediately 
fixed in 5% formaldehyde. In the laboratory the nematodes were removed from 
the sponges. In some cases the sponges were kept in sea water for three hours, in 
order to allow the nematodes to leave them. They were then fixed and mounted 
in glycerin following the Seinhorst method (1959). The coverglasses were sup- 
ported by splinters of broken coverglasses with a thickness of 0.11 mm. 


810 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


The specimens of Filipjev, Platonova and Sergeeva, and also L. bathybium, L. 
groenlandicum, L. caecum, and L. elongatum acephalatum had been mounted in 
glycerin-gelatin; for this study the three last-mentioned species were remounted. 
L. elongatum sensu de Man, 1893 and L. ranjhai had been remounted some years 
before. 

Specimens from Texel, used for the E.M.-study, were collected after they had 
left the sponge and were subsequently fixed in an iso-osmotic 1.5% glutaraldehyde 
solution buffered with sodium-cacodylate at pH 7.1 for 30 minutes. The head end 
was excised and embedded in 1% sea water agar. These agar pieces, measuring 
1 x 1 X 3 mm, were additionally fixed for one hour. Post-fixation took place in 
an 1% osmium tetroxide solution in 0.1 M sodium-cacodylate. 

After dehydration in ethanol, the material was transferred to monomere meth- 
acrylate in which it was kept overnight. The next day the monomer was replaced 
by pre-polymerized methacrylate, refreshened once and polymerized for 24 hours 
at 50°C. Sections were stained in uranyl acetate and lead citrate. 


Notation 


Cobb’s formula for expressing body proportions, which was used by Filipjev 
(1918), is of limited use for describing dimensions of populations, because no 
correlations can be given. In a hypothetical case where the length of individuals 
in a population varies from 6 to 9 mm, information is lost when the ratio ““b”’ is 
noted as 6.4—12.3. Moreover, the distribution remains indefinite. The standard 
deviation, which expresses the spread of the ratio, is useful only when applied to 
nematodes of equal length. The utility of the standard deviation is further de- 
creased, when applied to establish significant differences between Leptosomatum 
populations, by the fact that life cycle and environmental factors influence body 
length. 

To avoid indistinctness and to provide accurate information, body proportions 
are noted for each specimen separately in the Appendix. The specimens are ar- 
ranged according to body length to show the relation between body length and 
other dimensions. 

Body length was measured along the axis, which was drawn with the aid of a 
drawing-tube; the other, smaller measurements were taken directly with an ocular 
micrometer. Spicules were measured along the chord. The cephalic diameter was 
measured at the level of the cephalic sensilla, thickness of cuticle at level of the 
base of the pharynx. Pre-neural body length is distance from head end to the most 
anterior part of the nerve ring; length to ocelli and amphids are defined analo- 
gously. Body diameter at vulva level was measured when necessary, beside the 
protruding lips. The precision of the diameter of the amphid aperture is limited 
by focussing difficulties. 

Regarding the terminology, in this paper the term “‘lunula”’ is proposed for the 
crescent-shaped median lamella in the tail tip, surrounding the caudal pore as 
described by Hope (1967:313) for Pseudocella wieseri. The term “vaginal ov- 
ejector” is used to indicate the strong musculature in the vaginal wall of Deon- 
tostoma, Thoracostoma and Leptosomatides sp. which is depicted by Steiner 
(1916, Taf. 30 fig. 270, n) for what he considered to be Leptosomatum gracile 
(=Leptosomatides steineri Filipjev, 1922). 


VOLUME 96, NUMBER 4 811 


Artifacts 


A subject that has received little attention from taxonomists is the post-mortem 
phenomena caused by the fixative, the mounting medium or long-term storing. 
These phenomena may be advantageous—the fovea becomes more clear—but 
often they are disadvantageous, especially when not recognized: swelling of the 
cuticle, and dehydration after having been mounted for decades. 

Glycerin-gelatin shrinks when dried up as do the specimens mounted in this 
medium, and ruptures appear. This was the case in the type-material of L. e/on- 
gatum subsp. acephalatum Chitwood, 1936. On rehydration of glycerin-gelatin 
the medium increased in volume and ruptures disappear as a result of this swelling. 
The length of the nematode, which has been broken into pieces by the drying 
gelatin, increased by about 10%, which means that the original length has been 
restored. After removing superfluous gelatin, the specimen and adhering medium 
were dehydrated and mounted in glycerin in the usual way. Although the pieces 
of the nematode had not been measured, it seems acceptable that the length has 
been decreased by the same percentage as it increased by dehydration. This aspect 
of remounting is probably also applicable to the material of de Man (1893) because 
at present, these specimens are much smaller than originally described. 

I have also observed specimens, embedded in anhydrous glycerin for a consid- 
erable time, showing signs of shrinkage although they had been dehydrated suf- 
ficiently slowly to allow the glycerin to replace the water in the tissue. This 
phenomenon was noted in population 1-3 of L. bacillatum (pp. 820 and 821). 
The cuticula hardly changed but the pharynx and intestine decreased considerably 
in length, often resulting in a rupture in the intestine. The diameter of the body 
decreased: this can be seen quite readily when comparing the cephalic capsule, 
which hardly shrinks, with the more posterior tissues. This dehydration however, 
gives more contrast to the fovea of the male amphid. 

The type-material of L. ranjhai Timm, 1960 showed the same artifacts, but it 
is not known whether this is the result of remounting from glycerin-gelatin or of 
the above-mentioned factors combined. 

Body width is influenced by flattening more than the other dimensions. Often 
it was difficult to ascertain the degree of flattening, or even whether a specimen 
was flattened at all. For this reason body widths are considered of minor impor- 
tance. 

Purposely flattening in order to bring mounted specimens within focal distance 
of the immersion lens is to be avoided, especially when applied to type specimens. 
Glass rods or other supports for the coverglasses should have at least the same 
diameter as the nematode body. One holotype specimen studied had a body width 
of 130 um, whereas the supporting rods were no thicker than 24 and 28 um. 

In contrast, to soil-inhabiting nematodes, which are usually fixed after having 
actively passed through a cottonwool filter, marine nematodes are generally fixed 
together with the substrate. The fixed sample thus may contain specimens that 
were dead and decaying at the moment of fixation. It is, therefore, essential to be 
able to recognize post-mortem artifacts. For this purpose, nematodes that had 
died at least one day before, were fixed using 4% formaldehyde in sea water. 
Leptosomatum bacillatum showed the following artifacts: loosening and swelling 
of cuticular layers; loosing and retraction of the pharyngeal tissues at the anterior 
end. The cuticular pores became more distinct and the spicular manubrium be- 


812 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Table 1.—Differential characteristics of type-species of Leptosomatum and Leptosomatides. 


Cephalic capsule 


Sexual dimorphism in amphids 
Renette 


Vulvar glands in lateral hypodermal 
chord 


Intra-cuticular vulvar granula 
Vaginal ovejector 


Atrophy of digestive system and 
muscles in males 


Ventromedian precloacal papillae 


Specialized subventral pre- and 
postcloacal sensilla 


Spicules 
Gubernaculum 


Copulatory musculature 
Metanemes 


Leptosomatum elongatum 


reduced in male; poorly de- 
veloped in female. Poste- 
rior suture invisible 


present 
restricted to females 
absent 


absent 
absent 
present 


absent 
absent 


short and slender 


dorsal wall of spicule pouches 


slightly cuticularized 
not extensive 


loxometanemes 


Leptosomatides euxinus 


present; posterior su- 
ture visible 


absent 
wanting 


present 


present 
present 
absent 


present 
present 


robust 


complex; crura and cu- 
neus present 


strongly developed 


ortho- and loxometa- 
nemes 


came clearer. The turgor disappeared, resulting in longitudinal cuticular folds at 


both body ends. 


The Separation of Leptosomatum and Leptosomatides 


Leptosomatides euxinus Filipjev, 1918, and Leptosomatum elongatum Bastian, 


1865, are the type-species of their genera. Leptosomatum elongatum sensu de 
Man, 1893 is without doubt identical with L. elongatum Bastian, 1865. Both 
type-species have been studied and compared. The two genera can be distinguished 
by the characters listed in Table 1. 

One may wonder why Filipjev (1918) hesitated to transfer L-um arcticum? and 
L-um grebnickii, both described by him in 1916, to Leptosomatides. This may 
be explained in the following way. In 1912 Ssaweljev gave a poor description of 
a female, which read as follows: | 


“23. Leptosomatum tetrophthalmum n.sp. 2—12.7; a = 60; b = 7; c = 75. Der 
Bau des Kopfendes dhnlich wie bei Leptosomatum elongatum Bastian, 1865 
(de Man, 1893). Hinter den rotbraunen, kegelf6rmigen mit lichtbrechenden 
K6rperchen versehenen Augen noch ein Paar heller Pigmentflecke, 4hnlich wie 
bei den Enoplusarten. Nervenring am Ende des vorderen Oesophagusdrittels, 
Vulva am Ende des zweiten KGrperdrittels. Querfasernschicht der Cutis am 
Vorderende zu sehen. Palafjord, Mogilnojesee.”’ 


? [-um and L-ides are used in this section as abbreviations for Leptosomatum and Leptosomatides 
respectively. 


VOLUME 96, NUMBER 4 813 


No original material of Ssaweljev (1912) is present in the collection of the 
Zoological Institute in Leningrad but it is plausible that Filipjev saw this female 
(Platonova pers. comm.). In the collection, a slide is present (number 5267 dated 
12-IX-1915), from the same locality identified by Filipjev as L-um tetrophthal- 
mum Ssaweljev, 1912. Beside this specimen, some females are present, labelled 
L-um tetrophthalmum dated 22-IX-1925, and females, without additional eye 
pigment, labelled as L-um elongatum Bastian, 1865; both identified by Filipjev 
and published by Platonova (1967). 

All these specimens resemble L-ides euxines closely in the structure of the vulvar 
region. Assuming that the ovejector was characteristic for the type-species of 
Leptosomatum, Filipjev could not use it, to separate the two genera. 

Re-study of elongatum sensu Platonova, 1967 (sensu Filipjev), revealed that 
Filipjev was in error regarding the identity of L-uwm elongatum sensu Bastian, 
1865 and de Man, 1893. L-um elongatum sensu Filipjev and Platonova has all 
the characters diagnostic for females of the genus Leptosomatides as have L-um 
tetrophthalmum, L-um arcticum and L-um grebnickii. These characters are absent 
in L-um elongatum sensu de Man, 1893. 

This means that L-um tetrophthalmum sensu Platonova, 1967; L-um elon- 
gatum sensu Platonova, 1967 nec Bastian, 1865; L-um arcticum Filipjev, 1916 
nec Mawson, 1958 and L-um grebnickii Filipjev, 1916 belong to Leptosomatides. 
They will be discussed in another paper. L-um arcticum sensu Mawson, 1958 
will be discussed under L. kerguelense. 

No syntypes of L-um tetrophthalmum Ssaweljev, 1912, are present; it must be 
considered a species inquirenda. 


Morphological Observations 


In L. bacillatum, a cephalic capsule is present in females but not in males. De 
Man (1893) and Timm (1953) gave attention to this capsule in L. elongatum and 
the closely related L. acephalatum respectively. I will show that the reported 
difference between these two species, in structure of the cephalic capsule, does 
not really exist. 

The supposed difference in head structure between L. elongatum sensu de Man, 
1893 and L. elongatum subsp. acephalatum Chitwood, 1936 was the main reason 
for Timm (1953:230) to raise the latter to species level. For females of L. aceph- 
alatum Timm described “‘six pairs of fine sclerotized pieces, symmetrically ar- 
ranged around the ‘cap’ of oesophageal tissue,” which was presumed to be ho- 
mologous to “‘un systéme de deux lignes chitineuses et trés minces . . . et qui font 
défaut dans la region dorsale”’ as described by de Man (1893) in L. elongatum. 
This comparison is the result of an incorrect interpretation of de Man’s paper; 
neither de Man’s nor Timm’s passage concerns the cephalic capsule. 

De Man in fact described the anterior end of the ventrosublateral pharyngeal 
glands. He described the cephalic capsule as “‘une sorte de charpente chitineuse, 
radiairement symétrique et située a la péripherie, a laquelle s’insére évidemment 
Vextremité anterieure de l’oesophage.” 

In whole mounts, the cephalic capsules seems to be a refractive structure that 
quickly disappears out of focus and, therefore, Mawson (1958) described scler- 
otized pieces in what she considers to be L. arcticum and Timm (1960) described 


Fig. 1. Sections through cephalic capsule of female L. bacillatum at four different levels. A, Anterior 
part; B, Two microns posteriad of A; C, On level of cephalic papillae; D, Through posterior part of 
capsule. (CC: Cephalic capsule, LG: Ventrosublateral pharyngeal gland, DM: Dilator muscles, FO: 
Foramen, DS: Dorsal sector of pharynx.) 


VOLUME 96, NUMBER 4 815 


similar structures for L. ranjhai. Inglis (1964:289) is quite sure that what Timm 
(1953) is describing is the “‘lining of the cephalic ventricle, one component of 
each pair of sclerotized pieces corresponding to the musculature supplying the 
onchia and the other component corresponding to the radius of the oesophagus.” 

In Fig. 1A—D the structure of the anterior end ofa female is represented spatially. 
Section A is cut through the anterior part of the capsule; B two microns posteriorly; 
C at the level of the cephalic papillae and D just in front of the posterior suture. 

The cephalic capsule—the “‘charpente chitineuse”’ of de Man—is a conoid cap- 
sule; reduced but homologous to the capsule in Leptosomatides, Syringonomus, 
and the Thoracostomatinae. In pre-adult males this capsule is present; it disap- 
pears when the cuticle is shed during the last molt, so it is a part of the cuticle. 
In Fig. 1A—D this layer can be seen as an electron-dense layer that consists of 
radially arranged rods, 0.75 um in length. 

The posterior suture is almost straight; the anterior is interrupted by the inner 
labial sensilla, but neither suture is visible in glycerin-slides. De Man (1893) 
depicted this rim in his Fig. 9b. 

The anterior end of the pharynx is affixed to the cephalic capsule. Each sector 
of the pharynx contains four dilator muscles (Fig. 1 DM), paired two by two; 
these muscles were termed the “‘sclerotized pieces”’ by Timm (1953). The space 
between these bundles, the foramen (Inglis 1964), is filled by the socket cell of 
the labial sensilla on the inner and the pocket cell on the outer side; the latter, 
which is filled with electron-dense droplets, increases posteriorly in size and is 
pushed aside into the body-cavity at the posterior end of the cephalic capsule. 
The two paired bundles are separated by the pharyngeal nerves, apodemes and 
associated muscles, and ventrosublaterally by the pharyngeal glands. The cephalic 
ventricle (Inglis 1964) is absent in Leptosomatum. 

The “secondary capsule”’ as depicted by Filipjev (1916, Fig. 4a) is a space, filled 
with a spongy tissue, between the cuticular layers; I am not certain about its 
ultrastructure. This space might be homologous to the lunula. These secondary 
capsule and lunula have been underestimated as a diagnostic character in the 
Leptosomatidae. In males, if the cephalic capsule is lacking, this secondary capsule 
(Fig. 10b) may be confused with the cephalic capsule. 

In the anterior part of the pharynx I have observed one dorsal and two ven- 
trosublateral glands. The former empties into the pharyngeal lumen; the cuticu- 
larized duct is easily observed in glycerin specimens. The ventrosublateral glands 
(Fig. 1 LG) open on the lips as described by Timm (1953). These ducts are also 
cuticularized; de Man (1893) described them as “deux lignes chitineuses”’ being 
absent in the dorsal region. 

In contrast to the amphids in females and juveniles, the amphids in males are 
remarkable. In males the fovea is an inverted cardiform pouch with, in L. bacil- 
latum, a length of 10 um which opens to the exterior by a small pore. The fusus 
is about 15 um in diameter, fusiform, 40 um long, and leading to the amphidial 
gland (Fig. 10b). Some preliminary observations are worth mentioning. 

In L. bacillatum the amphidial glands are 600—900 um long and extend to the 
pharynx base. In related species with a short pharynx, the glands overlap the 
intestine. The posterior part is glandular and contains secretory organelles. The 
duct of the amphidial gland is filled with numerous microvilli (Fig. 2) with a 
diameter of 0.2-0.5 um: their number exceeds 500 in the posterior part of the 


816 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


eo sk ie 


Fig. 2. A, Section through posterior end of fusus in L. bacillatum, male. AG: Amphidial gland, 
DS: Dorsal sector of pharynx. B—D, Details of microvilli. (A, 10,000x; B, 100,000x; C and D, 
80,000 x.) 


fusus. Posteriorly the number decreases; at the level of the ganglia of the amphidial 
nerves, 50 could be counted. 

These microvilli are composed of alternating electron-dense and transparent 
layers, the latter on the outer side. The outer two or three electron-dense layers 
are circular without interruptions; interiorly these layers become irregular and 
single fibers are present in the center. In the anterior part of the fusus the villi 
are enclosed in membraneous chambers that resemble the amphidial chambers 
in the male of Meloidogyne incognita (see Baldwin and Hirschmann 1973). In 
the fovea 14 modified cilia could be counted. 

No features were observed that contradict the opinion that the villi originate 
in the anterior part of the fusus. This means that some of the villi reach a length 
of at least 400 um. However, as the free-floating ends of the microvilli are extended 
in the direction of the amphidial gland, it is difficult to imagine that, unless actively 


VOLUME 96, NUMBER 4 817 


involved in transport, the microvilli are not expelled by the excretory products 
of the amphidial glands. Numerous ganglia are situated anteriorly and posteriorly 
to the nerve ring in both sexes. In males two of them, at one corresponding body 
diameter behind the nerve ring, are swollen and therefore, I presume them to be 
the ganglia of the amphidial nerves (Fig. 9a). They are easily seen under a dissecting 
microscope. Under the light microscope the amphids in other species of Lepto- 
somatum resemble those in L. bacillatum. Bastian (1865) depicted these amphidial 
glands in the male of L. elongatum. 

The renette is situated ventrally in the posterior part of the pharynx and is 
restricted to females; in specimens with a short pharynx, ““b” exceeding 8, the 
gland partly or wholly overlaps the intestine. The renette is not always developed, 
but the pore is always visible in laterally mounted females. In males I have 
observed neither pore nor renette, and as hundreds of males have been studied, 
I am certain that they are absent. 

The renette in females might have been functionally replaced by the amphidial 
glands in males of Leptosomatum, although it will be difficult to prove this hy- 
pothesis. 

The ventromedian cells in the pseudocoelom, here termed coelomocytes to 
avoid the misleading term pseudo-coelomocytes, may be present in juveniles and 
adults. In some populations they are developed, in others not; populations occur 
in which these cells are restricted to a part of the population. Further details 
concerning their structure and function are wanting. 

Subventral pre- and postcloacal sensilla are designations for those specialized 
setae, often placed on hemispherical swellings of the cuticle, which differ from 
those in the subdorsal region, and which are functionally related to the role of 
the male, as depicted e.g., for Leptosomatides inocellatus by Platonova (1978:73). 
In L. punctatum, which is provided with cephalic setae, the setae are sparsely 
distributed over the whole body, just as other species with setae. They also occur 
in the subventral and subdorsal cloacal region. 

In the lateral epidermal chords of Cylicolaimus (see de Man 1889a:1) glands 
are present; in Pseudocella they are more simply built, and these glands are 
restricted to the vulvar region in Leptosomatides. These glands are absent in 
Leptosomatum as are the vaginal ovejector and intra-cuticular vulva granula, 
which, however, are present in Leptosomatides. 

The pre-cloacal ventromedian supplement is present in L. punctatum and in 
males of L. keiense. The copulatory musculature is not reduced in the former and 
a correlation may exist between the presence of the supplement and this mus- 
culature. 


Descriptive Section 


Leptosomatum Bastian, 1865 


Phanoglene Eberth, 1863 nec Nordmann, 1840 (Filipjev, 1918). 
Leptosomatum Bastian, 1865:144.—de Man, 1893:102-103.—Filipjev, 1918:42- 
44.—Platonova, 1976:58-—60. 


Type-species.—L. elongatum Bastian, 1865. 
Diagnosis.—Leptosomatinae Filipjev, 1918, with weakly developed cephalic 
capsule, apparent only in optical section; reduced in male or in both sexes. Somatic 


818 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


tissues atrophied in males. Renette and cervical pore restricted to females, inci- 
dentally present in juveniles. Renette usually situated in pharyngeal region, but 
not always developed. 

Sexual dimorphism expressed in structure of amphids. Males with enlarged 
fovea; amphidial glands strongly developed and outstretched over almost entire 
pharyngeal length. Pre- and postneural region of pharynx covered by numerous 
ganglia in both sexes. Ganglia of amphidial nerves in males much enlarged and 
situated at one body-diameter posterior to nerve ring. 

Stoma narrow, without onchia or odontia. Labial sensilla subcuticular, cephalic 
and cervical sensilla seti- or papilliform. Dorsal pharyngeal gland orifice at level 
of amphids in pharyngeal lumen; orifices of ventrosublateral glands on anterior 
end; ducts cuticularized. 

Ocelli provided with lens. Caudal glands long; overlapping intestine. Lunula 
present. Ventral row of coelomocytes usually present. Dorso- and ventrolateral 
orthometanemes present.* 

Male diorchic, testes opposed and outstretched. Female amphidelphic, anti- 
dromic. Gubernaculum simply built; without appendices. Precloacal ventrome- 
dian supplement reduced or absent. Subventral pre- and postcloacal genital sensilla 
absent. 


Leptosomatum abyssale Allgén, 1951 


Allgén described a female, originating from a depth of 400 m from the Sagami 
Sea near Japan, which was not available for this study. He mentioned the shape 
or the amphids—small and transversely oval—as different from L. elongatum. It 
would be interesting to know whether L. abyssale has ocelli. 

The description is absolutely inadequate; no details on the cephalic capsule, 
sensilla, or vulvar region are given. Until the slide is available for re-study, L 
abyssale must be considered a species inquirenda. 


The Leptosomatum bacillatum Complex 


To this complex belong L. bacillatum (Eberth, 1863), L. acephalatum Chit- 
wood, 1936, and L. sachalinense Platonova, 1978. These species might be con- 
specific, but in view of the geographical distribution and minor differences in size 
and ratios, I advise considering them as closely related species until well preserved 
material becomes available for a detailed comparison. 


Leptosomatum bacillatum (Eberth, 1863) Bastian, 1865 
Figs. 3-12 


Phanoglene bacillatum Eberth, 1863:19-20. 
L. elongatum Bastian, 1865:145. 

L. filipjevi Schuurmans Stekhoven, 1950:27. 
L. gracile Bastian, 1865:145-146. 

L. sabangense sensu Allgén, 1942:8. 

L. tuapsense Sergeeva, 1973:1710-1712. 

? L. sp. Kreis, 1928:139. 


3 For terminology see Lorenzen 1978. 


VOLUME 96, NUMBER 4 819 


Nec L. elongatum sensu Platonova, 1967; L. gracile sensu Allgén, 1954. (Both 
belong to Leptosomatides and will be discussed in another paper.) 


Diagnosis. —Cephalic and cervical sensilla papilliform. Cephalic capsule present 
in juveniles and females; absent in males. Ventromedian precloacal supplement 
absent. Caudal pore terminal. Ocelli relatively far posterior. Renette restricted to 
pharyngeal region. 

Distribution. — Mediterranean, Black Sea, North Sea, (Spitsbergen ?, Vancouver 
Island ?, California ?, South Georgia ?, Gulf of Panama ?, Argentina ?, Lesser 
Antilles ?, and Falkland Islands ?). 

I consider records with a question mark to be doubtful because of the numerous 
errors Allgén made in identifications of species of Leptosomatum and Leptoso- 
matides. For example, Leptosomatum microlaimum Allgén, 1957, is a species of 
Leptosomatides and has been transferred to that genus by Platonova (1976). 
Specimens identified by Allgén (1954) as Leptosomatum gracile are doubtful as 
he mentions the presence of vulvar glands, which are characteristic for Lepto- 
somatides. The specimens identified by Allgén as Leptosomatum sabangense be- 
long, as far as can be determined, to Leptosomatum bacillatum. Finally, it is 
doubtful that Allgén has accurately identified any of the species belonging to the 
Leptosomatum bacillatum-complex, given the morphological similarity among 
members of that complex and the superficial nature of Allgén’s work. 

Synonymy.—Eberth (1863:20) described L. bacillatum as Phanoglene bacilla- 
tum from: “‘unter Corallen im Hafen von Nizza.” Attempts were made to obtain 
material from the type-locality but harbor constructions had been carried out and 
in a letter dated 1980-1-22 Dr. A. Meinesz stated: “*. .. qu'il n’y a pas de ‘banc 
de coreaux’ dans le port de Nice et 11 n’y en a jamais eu.”’ Recently Marc Lavaleiye 
(pers. comm.) suggested that Eberth might have meant the calcareous alga Cor- 
allina. In (1878) de Man reported L. bacillatum from the Mediterranean, but as 
he did not make permanent mounts of the nematodes collected prior to 1876 
(Loof 1961), only the description can be used. 

Filipjev (1918) reported L. bacillatum from the Black Sea. These specimens 
are still present in the collection of the Zoological Institute in Leningrad where I 
was able to study them. Filipjev mentioned the presence of the opening of the 
gland of the accessory organ; this could not be confirmed. The amphids of the 
female were vaguely perceptible; presumably Filipjev depicted the male amphid 
in the figure of the female (Fig. la). In 1922 he reported gravid females with a 
length of 12.8 mm. 

Schuurmans Stekhoven (1950) renamed L. bacillatum sensu Filipjev, 1918, as 
L. filipjevi because Filipjev did not depict the cuticular pores on the tail tip. These 
pores are depicted by Eberth (1863) and were also present in the juvenile described 
by Schuurmans Stekhoven (1950). Examination of L. filipjevi Schuurmans Stek- 
hoven, 1950 (=L. bacillatum sensu Filipjev, 1918) showed that the pores are 
present. 

Leptosomatum elongatum Bastian, 1865, was described from Falmouth; this 
material has probably been lost. De Man (1893) gave a redescription based on 
specimens from the type-locality, and synonymized L. elongatum and L. gracile. 
These slides are still present in the collection of the Zoological Museum in Am- 
sterdam, and were placed at my disposal. They are labelled: 


820 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


A 57, Leptosomatum elongatum B. 2 Trefusis VI-1892. Zo6l. Museum A’dam. 
V. As. no. 652. 

A 58, Leptosomatum elongatum B. 6° Trefusis VI ‘92. Zod]. Museum A’dam. V. 
As. no. 653. 

A 58, Leptosomatum elongatum B. 6° Trefusis VI ‘92. Zo61. Museum A’dam. V. 
As. no. 654. 

A 59, Leptosomatum sp.? 6 Wimereux 1890. Zo6l. Museum A’dam. V. As. no. 
655. 


The nematodes were in a rather good condition; they only showed some shrinking 
caused by dehydration as mentioned before, but were identical to those recently 
found along the Dutch coast and L. bacillatum sensu Filipjev, 1918. 

Compared with the description, the mounted specimens of 1893 have decreased 
in size by approximately 30 percent. De Man was accustomed to studying spec- 
imens prior to mounting; only a part of his material was transferred to permanent 
slides (Loof 1961). Measurements were carried out on living specimens or on 
specimens recently fixed. It is known that an increase in the volume of a nematode 
in a hypo-osmotic environment, is expressed especially as an increase in body 
length. Thus de Man possibly measured his material in diluted seawater. Ac- 
cording to Newall (1976) the length of Enoplus brevis increases by 40% in a 10% 
diluted seawater solution. 

Leptosomatum tuapsense Sergeeva, 1973, was found to be identical to L. ba- 
cillatum. According to Sergeeva L. tuapsense differs from L. elongatum by the 
setae (?) and structure and length of the spicula. De Man (1893) mentioned a 
spiculum length of 98 wm, whereas Sergeeva gave 94 um; the length of the cephalic 
sensilla is 1.5 and 1.25 wm respectively. 

The holotype of L. tuapsense (slide N 8092), which is deposited in Leningrad, 
has been studied. The shape of the spicula, as depicted by Sergeeva, is not the 
lateral view; the manubrium is less cuticularized than depicted by her. The pres- 
ence of a cephalic capsule could not be confirmed; she depicted the ducts of the 
ventrosublateral pharyngeal glands. The dimensions of the amphids have to be 
halved; the breadth of the fovea is one-sixth of the corresponding body diameter. 

In the course of time, Sergeeva collected more material from the Black Sea, 
which was assigned to L. bacillatum (Eberth, 1863) and deposited at the Zoological 
Institute in Leningrad. I herewith synonymize L. tuapsense Sergeeva, 1973, with 
L. bacillatum (Eberth, 1863). 

From the Swedish Museum for Natural History, Stockholm, three slides were 
placed at my disposal. They are labelled: ““RMev Sthlm 3:13, 3:66, 3:82” and 
represent L. sabangense sensu Allgén, 1942:8. Although these juveniles are in a 
poor condition, I consider them identical to L. bacillatum. 

Regarding L. sp. Kreis, 1928, more information is desired. It might belong to 
L. bacillatum, the length and ratio “‘c’’ however, need confirmation. 


New. Records 


1. Den Helder, The Netherlands (52°58'N, 4°42’E); Nov 1970. 3 juv., 4 6 and 8 
2, collected from Polysiphonia sp. and Halichondria panicea. Littoral. Collec- 
tion Nematology Department Wageningen. 

2. Kattendijke, The Netherlands (51°33’N, 3°47’E); Oct 1970. 25 juv., 15 6 and 


VOLUME 96, NUMBER 4 821 


19 2, collected from Halichondria panicea. Littoral. Collection Nematology 
Department Wageningen. 

3. Burghsluis, The Netherlands (51°40'N, 3°40’E); Feb 1978. 200 specimens from 
Halichondria panicea. Littoral. Collection Nematology Department Wage- 
ningen. 

4. Texel, ‘t Horntje, The Netherlands (53°01'N, 4°47’E); Jun 1977. 230 specimens 
collected by Robin den Ottolander from Halichondria panicea. Littoral. Col- 
lection Nematology Department Wageningen. 

5. Texel, Oudeschild, The Netherlands (53°03’N, 4°50’E); Nov 1970. 55 juv. 10 
6 and | 2, collected from Halichondria panicea. Littoral. Collection Nematol- 
ogy Department Wageningen. 

6. Wimereux, France (50°48'N, 1°34’E); 1 2, collected by de Man in 1890 and 
labelled ““Leptosomatum sp.” Collection Zoological Museum Amsterdam. 

7. Ambleteuse, France (50°48’N, 1°34’E); Jun 1978. 2 2, collected by Michiel Buil 
from Halichondria panicea. Littoral. Collection Nematology Department Wa- 
geningen. 

8. Banyuls, France (42°29'N, 3°07’E); 3 2 from unknown sponges. Jun 1976. 
Collection Nematology Department Wageningen. 

9. N.E. England. | 2 and 1 6; collected by R. W. Warwick from Laminaria 
holdfasts at low tide on a rocky shore. Collection Smithsonian Institution 
Washington, D.C. 


Discussion and Description of New Records.— The general morphology has been 
described by de Man 1893 (L. elongatum), Filipjev 1918, and Timm 1953 (L. 
acephalatum). The ultrastructure of the cephalic capsule and amphids, has been 
described in a previous section. 

The study of the life cycle revealed an annual cycle for the Dutch population 
(4) and it is reasonable that this cycle is also present in other populations in 
temperate zones. The eggs are deposited in July and August; the length and 
development of the nematodes are correlated with the sampling date. There are 
indications that populations collected at corresponding days in different years 
show significant differences in length. This may be caused by food supply and/or 
temperature effects. 

The renette is maximally developed in autumn. In August 1978, more than 
50% of the females showed a more or less developed renette, whereas in the 
autumn of 1981 at the same locality (‘t Horntje) this gland was found in less than 
10% of the specimens. 

Pre-adult females of the February population from Burghsluis can have a body 
length between 4 and 6 mm; pre-adults of Texel (June) always exceed 6 mm in 
length, and these pre-adults reach a length of 8 mm. Schuurmans Stekhoven (1950) 
described a juvenile from Villefranche measuring 9.5 mm. 

Females continue growing after having reached the adult stage; for males, there 
is no evidence for length increase in the adult stage. The ratios are length-depen- 
dent and therefore correlated with the seasons. The ratios of the Dutch population 
are plotted on graphs to show the length dependance and _ variability 
(Figs. 3-6, 8). 

Newly hatched juveniles reach a length of 1.4 mm; the maximum length of 


822 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


OM160- 119 9+©6 e 
Sie!) ° 


0 1 2 3 4 5 6 7 8 9 Or at PA hi) 
Length (mm ) 


Fig. 3. Relation between ratio “‘a”’ (length/body width) and length, based on several L. bacillatum 
populations (Texel). 


females from populations 2 and 3 never exceeded 12 mm. Population 4 yielded 
adults of 14 mm whereas one of the females from Banyuls measured 16.8 mm. 
The variability of structure and length of spiculum and gubernaculum are given 
in Fig. 7. 
The ocelli of L. bacillatum are placed relatively far posteriorly compared with 


pe Sern eS 
2 69 Sateen $15) (i e 


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Lee eee 
* 5 OP ejeOrte 
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5 ones 
ie © (ete on c° 
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0 i 2 3 4 5 6 7/ 8 9 10 11 12 13 14 

Length (mm) 

Fig. 4. Relation beween ratio “b” (=length/pharynx length) and length, based on several L. ba- 
cillatum populations (Texel). 


VOLUME 96, NUMBER 4 823 


Me 180 


160 
140 
120 
100 
80 
60 
40 


20 


0 1 2 3 4, 5 6 7 8 9 10 11 V2 18) 14 


Length (mm ) 
Fig. 5. Relation between ratio “c” (length/tail length) and length, based on several L. bacillatum 
populations (Texel). 


L. kerguelense with which females could be confused. The distance from anterior 
end to ocelli, in the former species, is about 1.3 times the corresponding body 
diameter and up to 2.0 in the biggest females; in males it ranges from 1.4 to 1.7. 
In L. kerguelense the same calculation varies from 0.7 to 1.0 for females and 


VaBw Bosnia) ae 
Wd 


Length (mm ) 


Fig. 6. Relation between “‘tail/anal body width” and length, based on several L. bacillatum pop- 
ulations (Texel). 


824 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


} 


Fig. 7. Variability in spicule shape of the L. bacillatum population from Burghsluis. 


from 0.8 to 1.1 for males. Incidentally the ocelli lie at slightly different levels— 
one more anterior—but this phenomenon is not so common as in 
Leptosomatides sp. 

The transverse oval amphid aperture, often called ““amphid,” measures about 
1 um in females and is situated at 13-24 um from the anterior end; the opening 
leads to an almost round fovea with a diameter of 1.5—3.0 wm. The amphidial 
gland was never noted in females. In males, the canal through the cuticle is conical; 
the smaller anterior opening measures 1.0—1.5 um; posteriorly, at the level of the 
epiderm, this canal appears to be circular; 5 wm in diameter. The underlying fovea 


Vulva position N=84 
% 80 Burghsluis ° 
neXER ° 


5 6 7 8 9 10 11 12 13 14 
Length (mm) 


Fig. 8. Relation between V (=distance from anterior extremity to vulva x 100/body length) and 
length, based on several L. bacillatum populations from Texel. 


VOLUME 96, NUMBER 4 825 


is an inverse, obtuse cardiform pouch. The pore, especially in males, is difficult 
to resolve. 

The cephalic and cervical sensilla are papilloid, and reduced posterior to the 
nerve ring. The paired cephalic papillae are of different length; those situated more 
laterally reaching a length of 1.5—2 wm, the more medial papillae 1-1.5 wm. They 
are placed in cuticular invaginations and are in a number of cases difficult to 
detect. The papillae near the caudal gland pore are irregular in number and 
position. 

The cephalic capsule in females is refractive and attains a maximum length of 
8 wm. 


Leptosomatum acephalatum Chitwood, 1936 
(L. bacillatum-complex) 


Leptosomatum elongatum acephalatum Chitwood, 1936: 
L. elongatum.— Chitwood, 1951, nec Bastian, 1865. 


Diagnosis.—Same as L. bacillatum but lower a- and c-value. 

Distribution. — East coast USA (Beaufort, North Carolina) and Mexico (?). 

Discussion of status.—In 1936 Chitwood split off the variety L. elongatum 
acephalatum based on a male without cephalic capsule. In 1951 he united the 
variety with the nominate form after having found the female. Having studied 
material collected from the same sponge Hymeniacidon heliophila, Timm (1953) 
raised the variety to specific rank. 

Timm’s arguments were the difference in number of eggs, the fine suture around 
the head in de Man’s specimens, the lack of sclerotized pieces in the dorsal head 
region, and the sexual dimorphism in head structure and size. 

Regarding the number of eggs per female, in the Dutch population the number 
varies between zero and 55 and depends on the season. The uterus stretches with 
an increasing number of eggs. 

The fine suture around the head—the posterior suture of the cephalic capsule— 
was not noted in mounted specimens of the Dutch L. bacillatum populations, nor 
could it be detected in the specimens on which the 1893 description was based. 
In living or newly-fixed specimens I have sometimes noted this suture. 

Concerning the lack of sclerotized pieces in the dorsal region, a misunderstand- 
ing exists, which is discussed in a previous section. The sexual dimorphism, as 
expressed in the absence of a cephalic capsule in the male, was not described by 
de Man (1893), but as males were also mounted, it seems reasonable to suppose 
that he noted the dimorphism and considered the absence of the refractive capsule 
in the male as an artifact. 

The material of Chitwood (1936) (one male, USNM 33973) and Timm (1953) 
(one male, USNM 33986) has been re-studied; they are identical in structure to 
L. elongatum sensu de Man, 1893; to L. bacillatum sensu Filipjev, 1918, and the 
material from off the Dutch coast. Both males however, are in poor condition. It 
is not precluded that L. acephalatum is conspecific to L. bacillatum (Eberth, 1863) 
but regarding the difference in ratio c and the geographical distribution, the de- 
cision to synonymize these species is postponed till more material from the type- 
locality becomes available. 


826 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Fig. 9. L. bacillatum from Texel. A, Anterior end of male; B, Anterior end of female. 


VOLUME 96, NUMBER 4 827 


Fig. 10. JL. bacillatum. A, Head of female, 102; B, Head of male, 76101; C, Posterior end female, 
76102; D, Posterior end male, 76101; E, Vulvar region, (Burghsluis). 


828 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


WV 


100 UM 


Fig. 11. A and B, Spicules L. bacillatum sensu Filipjev, 1918; C, Spicules holotype L. tuapsense 


Sergeeva, 1973. 


= 


New Records 


. Quintana Roo, Mexico; North end of Ascension Bay; 4 2, 2 6 and 2 juv. 


Collected 7 Apr 1960 by F. C. Daiber and E. L. Bousfield at inlet behind Allen 
Pt. On mangrove roots; [sognomon alata, Melampus, sponges, amphipods, 


fiddler crabs, and anemones. Collection Smithsonian Institution, Washington, 
D.C. 


. Quintana Roo, Mexico; Allen Point, Ascension Bay. 13 Apr 1960. 1 9°, collected 


by W. L. Schmitt. Collection Smithsonian Institution, Washington, D.C. 


. Quintana Roo, Mexico; North end of Ascension Bay. 2 6, 4 2 and 1 juv. 15 


Apr 1960 by E. L. Bousfield and H. Rehder. Shore just east of Halfway Point. 
Turtle grass flats off the Point to sandy beaches and mangrove roots, sand 
varying from a very fine sandy-mud to a coarser shell sand. Collection Smith- 
sonian Institution, Washington, D.C. 


. Quintana Roo, Mexico; behind central part of Niccehabin Reef; 16 Apr 1960. 


Collected by W. L. Schmitt et a/. 1 juv. Collection Smithsonian Institution, 
Washington, D.C. 


. Quintana Roo, Mexico; Ascension Bay. Along shore near Suliman Pt. 17 Apr 


1960, W. L. Schmitt et a/. 1 juv. On rocks in littoral. Collection Smtihsonian 
Institution, Washington, D.C. 


. Quintana Roo, Mexico; South end Cozumel Island. North of Pta. Santa Maria. 


22 Apr 1960. E. L. Bousfield. 1 2, collected on shore. Collection Smithsonian 
Institution, Washington, D.C. 


Remarks.—The measurements of these specimens are given in the Appendix. 


In general, the specimens from Mexico deviate in slenderness and tail length from 
the Dutch L. bacillatum specimens. The diameter of the lens is 8 um, compared 
with 7 um in L. bacillatum. 


VOLUME 96, NUMBER 4 829 


\ é ) 
100 UM 


Fig. 12. Spicules. A, Holotype L. diversum; B, Paratype L. diversum; C, D and F, Paratypes L. 
sachalinense; E, Holotype L. sachalinense. 


Leptosomatum sachalinense Platonova, 1978 
(L. bacillatum-complex) 


L. diversum Platonova, 1978:495. 


Diagnosis.—Same as L. bacillatum, pharynx slightly shorter. 

Geographical distribution. —South Sakhalin. 

Leptosomatum diversum (lapsus diversus) and L. sachalinense (lapsus sacha- 
linensis) Platonova, 1978, were, according to the author, fixed in alcohol. This 
material shows, moreover, the characteristic artifacts of a post-mortem fixation, 
resulting in clearance of the cuticular “‘pores,”’ longitudinal folding of the body, 
clearing of the contours and swelling of the spicular manubrium. The alcohol 
caused the content of the lateral epidermal chord to dissolve as depicted for the 
anterior body part of L. diversum. This artifact is not restricted to L. diversum 
as one of the paratypes of L. sachalinense (slide 8013) shows the same phenom- 
enon. The posterior body parts of the specimens are folded, the cuticle separated 
irregularly and the precloacal papilla, as described and depicted, has to be ascribed 
to this. This “papilla” is situated on one of the subventral folds and could not be 
confirmed in the paratypes. In both species the cuticle is pierced by small pores; 
this is not a diagnostic character. The caudal glands, described as equal in length 
for L. sachalinense reach a length of 880, 1090, and 1350 um. A difference in 
head structure could not be noted; this structure is identical to that in L. bacil- 
latum. I herewith synonymize L. sachalinense and L. diversum. The holotype and 
the description of L. sachalinense closely agree. 


830 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


A 
50/um 
B 
100/um 
Fig. 13. JL. bathybium. A, Head; B, Posterior end. 


Ratio “b,”’ in the original description of L. sachalinense (females), has a higher 
maximum value than in the Dutch populations of L. bacillatum. However, per- 
sonal examination of the paratypes revealed that the maximum value of “b” (b = 
6.3—12.9) is not as high as indicated by Platonova, and in personal communication 
with her, it has been determined that the datum is in error. The only feature in 
which L. sachalinense differs from L. bacillatum (Eberth, 1863), is the structure 
of the spicula; in some cases the manubrium is open, in some cases closed and 
heavily cuticularized (or swollen?). Although I am not able to distinguish this 
species from L. bacillatum, because of the poor condition of the first, I postpone 
the synonymization until more material becomes available for re-study. 


Leptosomatum bathybium Allgén, 1954a 
Fig. 13a, b 


The holotype, a male from a depth of 4500 m was placed at my disposal by 
the Museum for Natural History at G6teborg. The type is labelled ““The Swedish 
Deep Sea Exped. Leptosomatum bathybium, Allgén.” 

Mounted on a slide, it could only be studied from one side. The fixation or 
way of mounting has caused a loss of contrast; only the cuticularized structures 
in cephalic and caudal region could be recognized. Moreover, the structures of 
the anterior region and their relative proportions could not be reconstructed due 
to flattening. 

The interpretation of the internal head structure does not conform to the de- 
scription given by Allgén (1954a). Each lip bears in the transverse plane a C- 
shaped cuticularized structure, the concave side medial. From the middle of these 
C-shaped structures, a connective piece runs to three “buccal rods” (or mandibles?) 
in the anterior part of the lumen. A cephalic capsule is present; the posterior 
suture of this capsule, which curves to produce shallow lobes, is distinguishable. 


VOLUME 96, NUMBER 4 831 


The diameter of the cyathiform amphid is 16 um, the aperture about 14 um. The 
cephalic setae are vaguely visible; probably 10 setae are present; 6 um in length. 
The pharyngeal part of the worm is twisted dorsally over 370°. This part is hyaline 
to such a degree that the nerve ring could not be found with certainty. With some 
reservation, it is located on 28% of the pharynx length. 

Ocelli are absent. The testis could not be located, and the caudal glands probably 
overlap the intestine. A ventromedian pre-cloacal sensillum is situated at 150 
um. In ventral view, the “papilla” is horseshoe-shaped with the open side pos- 
teriad. The lunula is absent. 

Concerning the reproductive system Allgén (1954a) states: ““Wegen der Lage 
des Tieres im Praparat was es leider sehr schwierig das Geschlecht des jungen 
Tieres zu bestimmen. Bei anwendung von Olimmersion habe ich doch im Hin- 
terabschnitt des k6rpers 2 Organe entdeckt, im welchen ich mit Vorbehalt die 
kurzen sehr diinnen am proximalen Ende angeschwollenen Spicula und das plumpe, 
unregelmaszig geformte akzessorische Stiick glaube gefunden zu haben.” 

Leptosomatum bathybium Allgén, 1954, must be considered a species inqui- 
renda. 

It may be necessary to erect a new genus for this species close to Platycomopsis 
but I prefer to await the urgently needed revision of the Leptosomatidae. 

In the collection of the Smithsonian Institution, Washington, D.C., some un- 
identified specimens are present from the Atlantic Ocean (4500 m) that might 
belong to the same genus. At present these slides are labelled “‘cf. L. bathybium” 
and are at the disposal of the next revisor. 


Leptosomatum behringicum Filipjev, 1916 


The material, on which the original description was based, was collected by 
Grebnickii in 1880 in the Bering Sea. Platonova (1976) gave a redescription in 
which she indicated two holotypes: 5780 and 5781. The first contains a complete 
female, the other a single head. Platonova and I have agreed to exclude 5781 and 
to designate 5780, deposited in the Zoological Institute, Leningrad, USSR, as 
lectotype. 

Filipjev (1916) remarked that the weak color of the eye pigment was caused by 
alcohol used as a fixative, and that older females show traces of disintegration. 

Specimen 5780 is mounted in glycerin-gelatin, situated dorsoventrally in such 
a way that the structure of vulva and vagina could not be determined; this difficulty 
was reinforced by the filled uteri. In the anterior gonad, 9 eggs are present; the 
posterior contains 14 eggs that are pressed against each other and have a flattened 
appearance. 

The type-specimen of L. behringicum which I have examined was in such poor 
condition that it was impossible to obtain additional information concerning the 
structure of head, vagina, vulva, and lateral epidermal chord. It is impossible, 
therefore, to decide whether it belongs to Leptosomatum or Leptosomatides, and 
it has to be considered a species inquirenda. 


Leptosomatum breviceps Platonova, 1967 


In 1967 Platonova described a nematode from Filipjev’s collection. The slide, 
numbered 7383, bears the superscription: VIII-1914, Barentz Sea, Kolski’j Golf, 


832 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


50 UM 


Fig. 14. Anterior end L. caecum Ditlevsen, 1923, Lectotype. 


I. Filipjev, L. brev. 2 uv.) and is deposited in the Zoological Institute, Leningrad, 
USSR. 

The description of 1967 and the redescription of 1976 did not mention that 
the description was based on a juvenile. The formation of the vulva had started, 
but was not completed. 

The cuticular layers were split, resulting in a space of 3.5 wm at the anterior 
end and 2 um at the pharyngeal base. This phenomenon might indicate that the 
last molt was imminent. According to Platonova’s (1967) illustration, the anterior 
end is orientated dorsoventrally; the posterior part laterally. Both illustrations are 
modified. I have found that neither the anal opening nor the rectum could be 
observed, the tail is probably longer than described. The anterior cervical setae 
are slightly shorter than the cephalic setae, and they are progressively smaller 
posteriorly. 

This specimen is extremely flattened, supported by glass-rods of 28 wm. The 
medial portion of the exteriormost cuticular layer is a more or less compact layer 
at the anterior end, comparable to the spongy layer in the male of L. bacillatum. 
This layer was incorrectly called ““head capsule’; the anterior part of the space 
was termed “‘stoma ring.’ The setae are broken or partly invisible. 

Having setiform cephalic sensilla, a ventrally orientated spinneret and no ce- 
phalic capsule, this juvenile resembles L. punctatum. As only one juvenile is 
known, I consider L. breviceps Platonova, 1967, a species inquirenda, the more 
so because it was found far outside the known area of L. punctatum. 


Leptosomatum caecum Ditlevsen, 1923 
Fig. 14 


In 1923, five specimens of a nematode were collected by Ph. Dollfus near 
Rockall Island from a depth of 240 meters and offered to Ditlevsen for exami- 
nation. Two slides were obtained from the Zoological Museum in Copenhagen, 
and Dr. Kirkegaard was so kind as to give permission to remount the nematodes, 
which had been embedded in glycerin-gelatin. One slide is labelled “Pourquoi 


VOLUME 96, NUMBER 4 833 


Pas SA 207 26 Prof. 240m. fond a Lophohelia. Dollfus. Leptosomatum caecum 
n.sp. Hj. Ditl.”; the other without indication n.sp. I have added 1342 and 1343 
respectively. The dimensions of the specimens (for the abbreviations see p. 852) 
were as follows: 


S SN L DNR PL CL NW PW MW AW V% 


F Saye SISO) Guh@ 1528 D0 ee) 104 SZ BO. CO 
Ja 34k Yoaa BIO 1340 85 8 82 a) 43) 850 6S 


Each specimen has little optical contrast, but slide 1343 shows, more or less, 
the contours of the cephalic capsule, which resembles that of Pseudocella and is 
herewith designated as lectotype; the other female, slide 1342, is too hyaline to 
observe the capsule. 

Ditlevsen (1923) incorrectly interpretated the position of the amphids; although 
rather hyaline, they are slightly perceptible and situated as usual in the lateral 
lacunae. I did not depict them. The cephalic setae, of which 10 are present, reach 
a length of 9-10 wm. The cervical setae do not exceed 6 um; their position on the 
left and right body halves is not alike. The lateral vulvar glands are present; the 
vaginal ovejector seems to be absent as are the pre- and postvulvar sensilla and 
groups of setae near the caudal pore. The caudal glands are short and restricted 
to the tail as depicted by Ditlevsen. In the lateral epidermal chord, big vacuoles 
or glands can be seen with a diameter of 40 wm. The cuticle thickness varies from 
6 wm at the pharyngeal base to 9 um near the anal opening. 

Although Filipjev probably did not examine these specimens, he suggests in a 
footnote (1927:94) that L. caecum might belong to Pseudocella, with which I 
agree. 

The transferring of L. caecum to Pseudocella makes P. caeca (Ditlevsen, 1923) 
a secondary homonym to P. coeca (Ssaeljev, 1912) according to art. 58 sub 1 of 
the Code. If not a synonym of one of the other nominal species in Pseudocella, 
L. caeca must be renamed; I propose to postpone this decision until a revision 
of Pseudocella. 


Leptosomatum clavatum Platonova, 1958 
Leptosomatum kerguelense Platonova, 1958:60—61, partim. 


Diagnosis.—Cephalic and cervical sensilla papilliform. Cephalic capsule in fe- 
male 10 um long. Ocelli far posterior. Ratio “c’’ less than in L. bacillatum of 
comparable size. Male unknown. 

Distribution. — Kerguelen and Macquarie islands. 

Discussion.—The identity of this species, which was found at the Kerguelen 
Islands, is fixed by the designation of the lectotype in 1968 (see discussion of L. 
kerguelense). Only females and juveniles are known. They differ from L. kergue- 
lense by the ocelli being situated far posterior at about 1.5 corresponding body 
diameters from the anterior end (in the type, which is severely flattened, the pre- 
ocellar length hardly exceeds the corresponding body diameter), a more slender 
body, longer tail (T/ABW = 1.5), and the caudal pore being situated terminally. 

Slides 5836, 6013, 7346, 7365, 7369, 7371, 7372 and 7377 belong to L. cla- 
vatum (lectotypte 5835). Because type-material of the Zoological Institute in Len- 
ingrad is not loaned, I was not able to measure the specimens in detail. The 


834 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Table 2.—Dimensions of L. clavatum. DF, distance to fovea; C, cuticle thickness at pharynx base. 
For other abbreviations see p. 852. 


Sn L DF DL Cc PL CL a b c V% Labelled as: 


5835 14,420 28 109 2 1909 187 67 7.6 77 60 = Allotype L. clavatum 
5836 12,430 29 118 8 1726 129 52 7.2 96 61 £Allotype L. kerguelense 
6013 10,860 24 126 10 1411 145 46 7.7 75 59 Paratype L. kerguelense 


measurements are presented in Table 2. Although slide 5835 is labelled as “al- 
lotype”’ it represents the lectotype because Platonova (1968) designated this slide 
as holotype. 


New record 


1. Macquarie Islands (54°32’S, 158°59’E); 15 Feb 1967. 3 juv., 11 2and 1 6; 112- 
124 m. Collection Smithsonian Institution, Washington, D.C. 


Remarks.—On morphological grounds, as far as is known, the population from 
the Macquarie Is. cannot be distinguished from L. bacillatum. It deviates by the 
body proportions 1.e., the placement of the ocelli, ratio “‘c’’ and, to a lesser degree, 
the body width. The cephalic capsule comes to 10 um, the amphidial aperture to 
3 wm, the fovea to 4 um, and the lens diameter varies from 6 to 9 um. 

In this population, mixed with L. kerguelense and L. sp. A (see p. 846), one male 
was present that also might belong to the latter. It resembles the male of L. 
bacillatum. The spiculum length is 78 wm, the gubernaculum 19 wm, and the lens 
diameter is 9 wm in dorsoventral view. The anterior part of the single male is 
twisted; dimensions of the amphids cannot be given. No figures are given since 
the females differ only in the above-mentioned characteristics. The redescription 
of the lectotype and information regarding the male are wanting. 


Leptosomatum groenlandicum Allgén, 1954 
Fig. 15 


The male specimen, on which the description was based, was placed at my 
disposal by the Swedish Museum of Natural History. It is labelled: RMev Sthlm. 
37.299 East Greenland King Osc.fj.N-37. The nematode, mounted in glycerin- 
gelatin, was remounted because air had penetrated under the coverglass. 

This male was curved in the shape of a “c.”” The length of 14.544 mm given 
by Allgén (1954), is the straight distance between the extremities. The length along 
the body axis came to 17.5 mm. The nerve ring is situated 580 wm from the 
anterior end; the lengths of pharynx and tail are 2950 and 270 um respectively. 
Ocelli are absent. The shortness of the gonads is remarkable; the anterior reaching 
a length of 478 um, the posterior 488 wm. The junction of these gonads is situated 
10.3 mm from the anterior end. The spicula are 160 um long; they are ensheathed 
by a gubernaculum that is characterized by a dorsal outgrowth with a membra- 
neous appearance. Ten cephalic setae are present; the lateral setae are broadened. 
Six rows of cervical setae are visible extending to the level of the nerve ring. 
Subdorsal of the cloacal aperture, 4 setae could be seen. The cuticle is thick, lunula 
absent, and the caudal glands could hardly be observed. 


VOLUME 96, NUMBER 4 835 


AC 
50um 


C 


Fig. 15. L. groenlandicum. A, Head; B, Posterior end; C, Gland in lateral epidermal chord at level 
of pharynx base. 


In the lateral epidermal chord, characteristic glands are present, which open to 
the exterior by a pore. These openings alternate dorsally and ventrally; sometimes 
2 im sequence open on the same side. The presence of a coffee-bean shaped 
structure as described by Lorenzen (1981:136) could not be confirmed. The am- 
phidial aperture is large and probably closed by a shield. Due to the flattening, 
the head structure is difficult to interpret; a cephalic capsule is present as are the 
cephalic ring and oesophageal capsule as termed by Inglis (1964). A tooth seems 
to be present in the pharyngeal lumen. More material however, is necessary for 
confirmation. 

This male resembles Leptosomatides inocellatus Platonova, 1967, which differs 
from other Leptosomatides species by the absence of ocelli, lateral vulvar glands, 
ovejector, and lunula; and by the presence of lateral epidermal glands and amphids 
comparable to those of Leptosomatum groenlandicum. A new genus must be 
erected for these two species. At present I prefer to consider Leptosomaium 
groenlandicum a species inquirenda. 


Leptosomatum indicum Stewart, 1914 


This species was found in September 1903 near Chilka Lake in India. Filipjev 
(1921), in the additional notes on his revision, reported that he could not obtain 


Fig. 16. L. keiense. A, Anterior end male; B, Posterior end male; C, Anterior end female; D, 
Posterior end female; E, Renette, 76047. 


VOLUME 96, NUMBER 4 837 


Stewart’s paper. Although the paper was available, I did not succeed in obtaining 
the specimen (Indian Museum no. ZEV 6142/7) on which the description was 
based. 

The female is mounted in “glycerin jelly-formalin”’ and the figures suggest a 
loss of contrast in the cephalic capsule. In the paper, the length of the nematode 
is not given but, calculated from the other data, must be 5 mm. The ocelli are 
situated at a distance of 0.56 mm from the anterior end; this seems questionable. 

Because of the cephalic capsule structure, L. indicum probably belongs to Deon- 
tostoma despite the fact that the nematode is rather small. But prior to transfer, 
re-study of this specimen and collection of new material is necessary. Until more 
specimens become available a more precise identification cannot be attempted 
and L. indicum Stewart, 1914, must be considered a species inquirenda. 


Leptosomatum keiense Micoletzky, 1930 
Fig. 16 

The original material could not be located. The following information is based 
on the description of Micoletzky (1930) and additional material from the Phil- 
ippines. 

Diagnosis.— Cephalic, cervical, and body sensilla setiform; cephalic capsule 
absent in juveniles and adults; ventromedian precloacal papilla reduced (or ab- 
sent?). Caudal pore ventral to terminus. Ocelli far posterior, provided with big 
lenses (10-11 um). Pharynx short. Renette may overlap intestine. 

Distribution.—Indonesia and Philippines. 


New records 


Philippines; collected by E. G. Menez from algae (Caulerpa and Eucheuma). 
Sublittoral. Collection Smithsonian Institution, Washington, D.C. 


Pangasinan; Telbang Cove (16°11'N, 120°03’E). 1 6, 1 2 and 1 juv. 3 Sep 1967. 
Pangasinan; Cangaluyan Is. (16°22’N, 120°00’E). 2 6, 1 9and 1 juv. 4 Sep 1967. 
Salcedo, Bolic Is. (11°05’N, 125°39’E). 2 juv. and 1 6. 10 Sep 1967. 

Great Santa Cruz Is. (5°52'N, 122°04’E). | juv. 18 Sep 1967. 

Zamboanga; Sacol Is. (5°56'N, 122°11’E). 1 juv. 19 Sep 1967. 


udot ate 


Additional description and discussion. —At first sight, this species resembles L. 
punctatum in the presence of setiform sensilla, the ventrally placed spinneret and 
the absence of a cephalic capsule. It can be distinguished by the greater diameter 
of the lens and the strong negative allometric growth of the pharynx. In some 
males the pre-cloacal papilla is absent. 

Leptosomatum keiense, as described by Micoletzky, is probably a species-com- 
plex. According to Micoletzky (1930:280): “das praanal gelegene Erganzungsorgan 
hatte ich nicht immer nachweisen k6nnen .. . die Augenlage ist veranderlich .. . 
Linsendurchmesser 5.8—6.7, selten bis 12 um... die Ausbildung des Kopfborsten 
zeigt individuelle Verschiedenheiten ... b = 4.5-12.1.” 

The material from the Philippines shows the same variability. Males 76044 
and 76045 (pop. 2), which are mounted in lateral position, do not show any trace 
of an accessory organ; in male 76041, mounted dorsoventrally, a papilla is present 
141 wm anterior to the cloacal vent, whereas in 76049, mounted laterally, this 
papilla lies at 126 um. Moreover, female 76040 (pop. 1) seems to be provided 


838 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


with a ventromedian sensillum; this observation could not be confirmed in the 
other female (76046, pop. 2), which is twisted. The diameter of the lens is variable; 
7-13 um. Male 76049 (pop. 3) has a lens of 13 wm diameter, the two males of 
population 2 have a lens diameter of 7 um, but in general it varies between 9 and 
11 um. 

Notable is the short pharynx in one of the males (b = 11). According to Mi- 
coletzky (1930), this phenomenon also occurs in females. In juvenile 76048, a 
renette is developed that extends posteriorly 700 um from the anterior end and 
overlaps the intestine. The amphidial glands in 76041, 76033, 76045 and 76049 
attain lengths of 640, 800, 710 and 740 um respectively; in the latter the glands 
overlap the intestine. 

The ocelli, situated at 1.5 times the corresponding body diameter from the 
anterior end, are provided with an intensively pigmented cup. The cephalic setae 
are 3.5—4.5 um long; short setae are situated over the whole body length. The 
amphids, 15—27 um from the anterior end, resemble those of related species. The 
fovea measures 8 um in the male and 3 um in the female; the apertures 2 and 1 
um respectively. In one of the females (76040) the cervical pore could be detected 
at 227 wm from the anterior end. The spiculum length is 63-66 um, the guber- 
naculum, if present, was not perceptible. Male 76045 is extensively atrophied; 
numerous coelomocytes of 6 X 4 um could be seen throughout the body length 
of this male. 

As stated above, L. keiense might be a mixture of at least two species. The ratio 
““pharnyx length/tail length’? (=P/T) clearly separates the adults of population 2, 
in which P/T exceeds 12, from the remaining specimens in which P/T never 
exceeds 10. It is possible that the material of Micoletzky (1930) is still present 
and since the few specimens from the Philippines were collected at different 
localities, I postpone the separation, but I give some remarks regarding the spec- 
imens of population 2. 

These males can be distinguished by their relatively small lenses (7 um), the 
absence of a precloacal papilla, a slender body (‘‘a” = 97 and 104), a short tail 
(‘c?? = 94 and 104), and the allometric trend less conspicuous than the other 
males. Ratio “c’’ of the female, that has been found in the same sample, is also 
rather high; the lens diameter is 12 wm in dorsoventral view. The juveniles from 
population 2 cannot be distinguished from those of the other localities. 


Leptosomatum kerguelense Platonova, 1958 
Figs. 17a, b, 18 


Leptosomatum crassicutis Platonova, 1958:12—13 
Leptosomatum clavatum Platonova, 1958:15-16 partim. 
Leptosomatum arcticum sensu Mawson, 1965:315-316 


Diagnosis.— Cephalic and cervical sensilla papilliform; cephalic capsule present 
in both sexes; ventromedian precloacal papillae absent. Caudal pore slightly 
ventral to terminus. Ocelli relatively far anterior. Renette restricted to pharyngeal 
region. Tail length equal to anal body width. Spicula long; gubernaculum reduced 
to 2 membranes. 

Type.— Lectotype 5833; Zoological Institute Leningrad. 


VOLUME 96, NUMBER 4 839 


BCD 
100/um 


AE 
50 jum 


Fig. 17. Leptosomatum kerguelense (76056): A, Head; B, Caudal end. (C-E) L. punctatum: C, 
Caudal end of male, 1286-6; D, Caudal end of female, 1286-5: E, Anterior end of female, 76037. 


840 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


100 UM 


Fig. 18. Spicules L. kerguelense. A, Holotype; B, Paratype. 


Distribution. —A subantarctic species: South Georgia, Kerguelen Islands, Heard 
Island, Crozet Islands, and Macquarie Islands. 

Synonymy.—The three species from the Kerguelen Islands (Platonova 1958) 
were described from the remainder of an alcohol collection from which the mac- 
rofauna was removed. This fixation caused some particular effects of which the 
swelling and loosening of the cuticle are the most striking. The specimens are 
mounted in glycerin-gelatin, flattened, and as stated by Platonova (1958) in me- 
diocre condition. In 1968 the species were redescribed, depicted (some shifting 
had taken place) and holotypes (=lectotypes) were designated. 

Both papers are rather confusing; regarding L. clavatum for example, the lec- 
totype (slide 5835) is called holotype and labelled as allotype. Slide 5835 does 
not correspond with the formula: the ratios as given in the description are also 
at variance with both the formula and the type. The lectotype fits neither the 
description nor the figures. The male on slide 7633, labelled as holotype, does 
not belong to the syntype and more juveniles are designated as paratypes than 
originally belonged to the syntypes. 

In the description of L. crassicutis attention is given to the aberrant construction 
of the cephalic capsule. This feature, however, is an artifact due to the swelling 
of the cuticle, which can be confirmed by observing the cuticular pores, forming 
little holes in the cuticle surface and cones on the epidermis. 

The cuticle thickness is stated to be a differentiating character for L. crassicutis. 
I have measured the cuticle at the level of the pharyngeal-intestinal junction, as 
did Platonova, and the pre-ocellar body length. The data are given in Table 3. 
The cuticle thickness, which is heavily influenced by the fixative (or post-mortem 
fixation?), cannot be maintained as a diagnostic character for L. crassicutis. The 


VOLUME 96, NUMBER 4 84] 


Table 3.—Sex (S) or stage, Slide number (SN), Pre-ocellar body length (OL), Cuticula thickness 
(Cut.) and species assigned to Kerguelen population described by Platonova (1958). 


S SN OL Cut. Spec. S SN OL Cut. Spec. 
M 5831 75 12,1 crass J 7346 89 4,4 kerg 
F 5832 62 8,5 crass J 7359 69 8,4 kerg 
M 5833 73 8,5 kerg. F 7362 65 3,2 clav 
F 5834 89 6,1 kerg. J 7363 32 2,4 clav 
F 5835 109 2,0 clav. J 7364 65 1,6 clav 
F 5836 118 8,5 kerg. J 7365 97 4,0 clav 
F 5838 73 12,1 clav. J 7366 65 5,3 clav 
F 5840 69 2,4 clav. J 7367 32 2,0 clav 
M 6010 69 7,7 kerg. J 7368 57 5,7 clav 
M 6011 73 12,1 kerg. J 7369 105 5,7 clav. 
F 6012 77 12,6 kerg. J 7370 69 4,0 clav 
F 6013 126 10,1 kerg. J 7371 93 3,6 clav 
M 6014 77 8,9 kerg. J 7372 105 x clav. 
F 6015 V7 8,1 kerg. J 7374 57 4,0 clav 
F 6019 69 4,9 kerg. J 7377 81 3,2 clav 
M 7343 77 13,0 kerg. J 7633 57 8,1 clav 
J 7345 57 4,9 kerg. M 7634 64 2,0 clav 


lack of bristles on the surface of the cuticle is given as a distinguishing feature for 
all three species. Herewith I synonymize L. crassicutis and L. kerguelense as their 
lectotypes 5831 and 5833 are evidently conspecific. I propose the name L. ker- 
guelense Platonova, 1958; Recommendation 24a is not followed because the 
description of L. crassicutis, which is based on artifacts, would only lead to 
confusion. The type (5833) corresponds with the formula and description; the 
figure, however, is of one of the paratypes as the lectotype is mounted dorso- 
ventrally. The spicula of holo- and paratype are depicted in Fig. 18. The caudal 
glands, as depicted for L. kerguelense by Platonova, are much longer; they overlap 
the intestine as is usual in Leptosomatum. 

Measuring the syntypes, another feature was found that was not previously 
recorded. Two types of juveniles and females occur; the first belongs to L. ker- 
guelense, the other resembles L. bacillatum. These latter specimens are charac- 
terized by, among other features, ocelli situated far posteriorly. To this species 
belong slides 7346 and 5835, both labelled L. clavatum and slides 7365, 7369, 
7371, 7372, 7377, 6013 and 5836 labelled L. kerguelense. Slide 5835 is the 
lectotype of L. clavatum, and 5836 the lecto-allotype of L. kerguelense. The 
lectotype of L. clavatum does not agree in every respect with the description of 
1958. The measurements of the lectotype and discussion of its status are given 
in the paragraph of L. clavatum. 

Leptosomatum arcticum sensu Mawson, 1958; nec Filipjev, 1916 is also inden- 
tical to L. kerguelense. Mawson’s material was not available for this study. Based 
on the description, there is no need to assign this species to Leptosomatides because 
structure of the gubernaculum is as in other species of Leptosomatum. The only 
difference from L. kerguelense is that the spiculum/gubernaculum ratio is not 
identical. This might be caused by artifacts or the gubernaculum may be obscured 
by the opaqueness of the surrounding tissue. 


842 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


New records 


1. South Georgia (53°52’S, 37°37'W). 3 6, 2 2, 1 juv. Coll. 7 Feb 1966 at a depth 
of 97-101 m. Collection Smithsonian Institution, Washington, D.C. 

2. Macquarie Islands (54°32'S, 158°59’E). 5 Feb 1967. One pre-adult 2. 112-124 
m. Collection Smithsonian Institution, Washington, D.C. 


The above-mentioned specimens of the first population are in a poor condition 
which may be caused by a post-mortem fixation, but they clearly belong to L. 
kerguelense. The cephalic capsule in male and female measure 4 and 6 um re- 
spectively. The amphidial fovea has a diameter of 5 wm in the female and 11 wm 
in the male. The construction of these amphids is identical to that in L. bacillatum,; 
only slightly more robust. The tail is obtuse, caudal glands are long, and the caudal 
pore is shifted ventrally. Although this species is easily recognizable, a redescrip- 
tion from well preserved material is desirable. 


Leptosomatum micoletzkyi Inglis, 1971 


Remarks.—This species, described from one male, is distinguished from the 
species of Leptosomatum by the amphids, which lie at more than one cephalic 
diameter from the anterior end; the presence of subventral precloacal setae, and 
the absence of the lunula (?). Sexual dimorphism, comparable with Leptosomatum, 
seems to be absent. The spicules are slightly sinuous and end distally in blunt 
tips. The gubernaculum enfolds the spicules near their distal ends; proximally it 
forms large membranes. 

The male, which was not available, does not fit any nominal genus. At present 
it would lead to confusion to erect a new genus for this species and I consider L. 
micoletzkyi Inglis, 1971, a species incertae sedis until the female is described. 


Leptosomatum pedroense Allgén, 1947 


Allgén (1947) described this species from a juvenile which was not available 
for this study. The length of this juvenile is 7120 um with a ratio “a” of 29.06. 
This means that the diameter of this specimen is 240 um, leading to the assumption 
that this juvenile is extremely flattened. This is supported by the figure of the tail. 
Being based on a juvenile and described insufficiently, L. pedroense Allgén, 1947, 
must be considered a species inquirenda. 


Leptosomatum punctatum (Eberth, 1863) Bastian, 1865 
Fig. 17c-e 


Phanoglene punctata Eberth, 1863:20. 
? Stenolaimus macrosoma Marion, 1870:17-18.—1870a:10. 
Leptosomatum longisetosum Schuurmans Stekhoven, 1943a:4. 


Diagnosis.— Cephalic, cervical and body sensilla setiform; cephalic capsule ab- 
sent in juveniles and adults. Ventromedian precloacal supplement present. Cau- 
dal pore ventral to terminus. Ocelli relatively far posterior. Lens 6—7 wm. Cop- 
ulatory musculature relatively strongly developed. 

Distribution. —Mediterranean, Black Sea, and Red Sea. 

Synonymy.—Filipjev (1918) synonymized S. macrosoma Marion, 1870, with 
L. bacillatum (Eberth, 1863) because of Marion’s statement (1870:17), “Elle ne 


VOLUME 96, NUMBER 4 843 


presente pas non plus la couronne de soies longes et robustes”’ in which opinion 
he was followed by Platonova (1976). 

The argument of Filipjev, however, was based on an incorrect interpretation 
because Marion’s (1870:17) passage had been taken out of context. The whole 
paragraph reads: “Le tube oesophagien se termine en effet de la méme maniére 
que celui du Stenolaimus lepturus, mais la téte réguliérement arrondie ne porte 
point de papilles. Elle ne presente pas non plus la couronne de soies longues et 
robustes da sa congénére; la peine si l’on remarque quelques poils courts dispersés 
tout le long du corps et un peu plus nombreux 4a la téte.”’ In other words, S. 
macrosoma has neither papillae on its head, nor the long hairs of the proceeding 
species S. lepturus (=Anticoma acuminata (Eberth, 1863) op. Allgén, 1942). The 
latter has three protruding lips to which Marion alluded and called papillae. 

Stenolaimus macrosoma was characterized (Marion, 1870:17) by: “‘quelques 
poils courts dispersés tout le long du corps et un peu plus nombreux 4a la téte.”’ 
Therefore, it is obvious that S. macrosoma is more closely related to L. punctatum, 
as the sensilla are setiform. I believe S. macrosoma to be identical to L. punctatum 
(Eberth, 1863). 

In 1943a Schuurmans Stekhoven described L. /ongisetosum and, in the same 
year, illustrated it in a separate paper (1943b). I was not able to locate this 
specimen. According to the author, L. /ongisetosum differs from L. punctatum in 
the shape of the tail. This tail however, shows the typical shape of a juvenile of 
L. punctatum. 1 consider L. longisetosum identical with the latter. 


New records 


1. Red Sea; Ain Sukhna (29°36'N, 32°24’E). 1 2. Collected by W. D. Hope on 5 
Jan 1967 from the intertidal sediment on the beach at El Sokhna Hotel. Various 
types of corals and colonial coelenterates. Collection Smithsonian Institution, 
Washington, D.C. 

2. Banyuls; France (43°00, 3°10’E). 1 juv., 1 2and 1 6. Deposit of unknown sponges 
and Corallina. Jun 1976. Collection Nematology Department Wageningen. 


Description and discussion.—In general this species is smaller than L. bacilla- 
tum. The female from the Red Sea is only 3.6 mm long; half as long as the adults 
from Banyuls. 

The length of the cephalic setae varies among individuals. Filipjev (1918) men- 
tioned 6 um; longer than in this material where 3—4 wm has been measured for 
the juveniles, 4, 5 for the male and 4 and 5 um for the females. Setae of 2-3 um 
are sparsely present over the whole body length. Dr. Platonova was so kind as to 
remeasure the cephalic setae of Filipjev’s material, and found 3.5 um for the males 
and 4.9 wm for the females. 

The amphidial aperture and fovea in the male measure | and 8 um against 2 
and 3 in the female. The structure of these amphids is comparable to those in L. 
bacillatum. Eberth (1863), in his original description, probably depicted one of 
the amphidial glands. 

The ocelli, with a lens diameter of 6—7 um, lie at 1.3 times the corresponding 
body width from the anterior end. The spiculum and gubernaculum length are 
79 and 20 um, respectively. The ventromedian pre-cloacal papilla is situated at 
166 um from the cloacal aperture. The tail, in both sexes, is rather long, 1.8 times 
the anal body width and more tapered than in L. bacillatum. 


844 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Leptosomatum ranjhai Timm, 1960 


From the Arabian Sea Timm (1960) described a new species without sexual 
dimorphism in the head structure. The holo- and allotype were present in Wag- 
eningen in 1973 when a fire destroyed the laboratory and damaged a part of the 
collection. The types of L. ranjhai have been remounted but the contrast was 
almost lost which may be partly due to clearing in lactophenol. 

The cephalic capsule in L. ranjhai is strongly developed; the posterior suture 
was clearly visible but the presence of cuticularized rods, surrounding the head, 
could not be confirmed. 

The lateral epidermal chord in both sexes contains big vacuoles or glands, the 
structure of which could not be clarified; they measure from 22 x 19 to 34 x 30 
um and more than 30 could be counted in one body side. Ortho- and loxo- 
metanemes-I are present. A sexual dimorphism in the amphids is absent; the 
structure of vagina and vulva preclude placement in Leptosomatides. It is nec- 
essary to erect a new genus for this species. This will be done after having compared 
the other genera in the Leptosomatidae and the remaining material of L. ranjhai. 


Leptosomatum sabangense Steiner, 1915 
Fig. 19c, d 


Nec L. sabangense sensu Micoletzkyi, 1930 (=L. sundaense new name). 


Diagnosis.— Cephalic and cervical sensillae papilliform. Cephalic capsule pres- 
ent, posterior to cephalic sensilla. Tail length twice anal body width. Caudal pore 
terminal. Male unknown. 

Distribution.— Indonesia and Red Sea. 

Synonymy.—In 1915 Steiner described L. elongatum var. sabangense, which 
was raised to species level by Filipjev in 1921. Steiner split off the variety because 
of the tail length which is twice the anal body diameter. 

The figures of Steiner (1915) give another characteristic in which L. sabangense 
differs from L. elongatum, namely that the cephalic capsule is situated posterior 
to the cephalic papillae. This is depicted in both Figures 5 and 6 (Taf. 22) and 
confirmed in the text. This phenomenon is unique in this genus and needs con- 
firmation. 

Leptosomatum sabangense sensu Micoletzky, 1930, has another type of capsule, 
situated as usual in this genus but longer (11-13 wm). The pharynx is variable in 
length (b = 6.8—12.6). In the female, the ocelli are situated far posterior, in the 
male at less than one corresponding body diameter from the anterior end. More 
information is necessary concerning L. sabangense sensu Micoletzky, 1930 nec 
Steiner, 1915 which has to be renamed. I propose the name L. sundaense new 
name. 


New record 


1. Red Sea; Ain Sukhna (29°36’N, 32°24’E). 1 juv. Collected by W. D. Hope 5 
Jan 1967 from the intertidal sediment on the beach at El Sokhna Hotel. Various 
types of corals and colonial coelenterates. Collection Smithsonian Institution, 
Washington, D.C. 


Description.—This juvenile is characterized by the placement of the cephalic 
capsule posterior to the cephalic papillae and the tail length, which is 2.16 times 


VOLUME 96, NUMBER 4 845 


os ney 


Fig. 19. Anterior end 4, and head B, of L. sp. A; Tail C, and anterior and D, of L. sabangense, 
juvenile. 


the anal body width. The measurements are given in the Appendix. Tail and 
cephalic end are depicted in Fig. 19c, d. 
Discussion.—Records of L. sabangense which are only based on ratio c are 


846 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


doubtful because a correlation is necessary between this length and the anal body 
width resulting in a T/ABW of 2. Moreover in newly hatched juveniles this ratio 
may exceed 2.5 in L. bacillatum too. So a record of L. sabangense must be based 
on the T/ABW of adults, ratio c, and the placement of the cephalic capsule. L. 
sabangense sensu Micoletzky, 1924, a long-tailed female, also from the Red Sea, 
may be related to the above described juvenile. The record of Allgén (1942) is 
discussed under L. bacillatum. 

Despite the fact that at a juvenile is assigned to L. sabangense Steiner, 1915, 
in this paper, more information is necessary concerning this species, which must 
be considered a species inquirenda. 


Leptosomatum sp. A 
Fig. 19a, b 


Material. — Macquarie Islands (54°32’S, 158°59’E); 15 Feb 1967. 3 juv. and 6 
2; 112-124 m. Collection Smithsonian Institution, Washington, D.C. 

Description. This species, of which only juveniles and females are known, re- 
sembles L. bacillatum (Eberth, 1863) in the presence ofa cephalic capsule, cephalic 
papillae and the terminal caudal pore. It can be distinguished by the short pharynx 
and related to this, the renette in the postpharyngeal region, the short cephalic 
capsule, large lens, and slenderness of the anterior body region. The male (76081) 
assigned to L. clavatum might eventually be assigned to this species. 

The pharynx in the adults is relatively short, resulting in a ratio “b” of 9.8- 
11.8. The renette, which seems to be bilobed is developed in 76067, 76069, 76071, 
and 76072, and restricted to the anterior-intestinal region. The pore could not 
be observed. 

The cephalic capsule is relatively short, 6—8 um in length; as in the other species 
of this genus the posterior suture is not perceptible. The amphidial aperture and 
fovea measure 2 and 3 um respectively. The ocelli lie at 70-86 um from the 
anterior end, the lens varies from 8 to 10 um. The anterior body end is very 
slender; the cephalic width never exceeds 30 um, the width at the ocelli never 
exceeds 60 um. 

Discussion.—This species which appears to be hitherto undescribed, is not 
named because males are absent. Regarding the position of the renette I doubt 
whether this species belongs to Leptosomatum. An unpublished scanning study 
of the head of 504 and 518 however, revealed that the labial region is identical 
to that in Leptosomatum. Until more specimens become available, a diagnosis 
of this species cannot be given. 


Syringonomus Hope and Murphy, 1969 


The collection which was made available by the Smithsonian comprises 4 
juveniles and 4 females of Syringonomus typicus Hope and Murphy, 1969. The 
specimens were collected 20 Feb 1967 at a depth of 943-1007 m near Recife 
(7°58.0’S, 34°17.0'W). This species will be discussed here as this monotypic genus 
is Closely related to Leptosomatum. 

Dr. W. D. Hope was so kind as to send me a male and female paratype; 
comparison confirmed the identification of this deep-sea species. The measure- 
ments are presented in the Appendix. Little needs to be added to the description 


VOLUME 96, NUMBER 4 847 


of Hope and Murphy (1969). I wish to consider four points: (1) At the posterior 
end of the pharynx, in the male paratype, the amphidial glands are visible. These 
glands are easily overlooked as the males are not atrophied. (2) The renette is 
probably sexlinked; it is present in some females and absent in the male. (3) The 
subventral precloacal papillae are not specialized, they are comparable to the 
subdorsal setae. (4) Dorso- and ventrolateral orthometanemes are present. 

This genus is characterized by the unique lyre-shaped pattern on, and thickening 
of, the cuticle in males at the level of the amphidial aperture, and the absence of 
ocelli. Females are distinguishable from Leptosomatum species by the absence of 
ocelli and combination of cephalic setae and presence of cephalic capsule. 


General Discussion 


The genus Leptosomatum formerly comprised all species of Leptosomatidae 
with a reduced cephalic capsule, but now it is one of the most distinctly demarcated 
genera within the family. Together Syringonomus and Leptosomatum form a 
taxon that may be regarded holophyletic, just as each genus is in itself holophyletic. 

The presence of vaginal ovejector and lateral vulvar glands is a good character 
to distinguish females of Leptosomatides from those of the above-mentioned 
genera. The ovejector has been underestimated as a diagnostic character and may 
serve to separate Pseudocella, in which the ovejector is absent, from Thoracostoma 
and Deontostoma species. The same applies to the glands in the lateral epidermal 
chord, present in Pseudocella but restricted to the vulvar region in Leptosomatides, 
Thoracostoma, and Deontostoma as far as is known. In this way Thoracostoma 
species without ocelli can be distinguished from Pseudocella species if only females 
are at hand. In a separate paper I shall consider this in more detail and examine 
the systematic consequences. 

Reviewing these characters, Leptosomatides shares more characters with 7hor- 
acostoma and Deontostoma than does Pseudocella. Leptosomatum ranjhai Timm, 
1960, is closely related to Pseudocella; in L. ranjhai, lateral epidermal glands are 
also present, the pigment spots are situated anteriorly, the vaginal ovejector is 
absent and moreover, L. ranjhai is provided with loxometanemes, as are Pseu- 
docella, Thoracostoma, and Deontostoma as far as is known. Although metanemes 
are often difficult to observe, I am confident that loxometanemes are absent in 
Leptosomatum and Syringonomus. 

Hitherto I have been unable to study Paraleptosomatides Mawson, 1956; judg- 
ing from literature, this genus is related to the members of the Thoracostomatinae. 
The type-specimen of Leptosomella acrocerca Filipjev, 1927, has been lost (Pla- 
tonova pers. comm.), and according to Hope (pers. comm.) the type-material of 
Tubolaimella is also lost. 

The present author is still interested in re-studying the above-mentioned genera 
as well as Leptosomatum abyssale Allgén, 1951; L. indicum Stewart, 1914; L. 
keiense Micoletzky, 1930; L. micoletzkyi Inglis, 1971; L. pedroense Allgén, 1947; 
L. sabangense Steiner, 1915; L. sabangense sensu Micoletzky, 1930, and L. ba- 
cillatum, L. elongatum, L. gracile and L. sabangense sensu Allgén as described 
in several papers. 

As no syntypes are present, attempts will be made to obtain material to designate 
neotypes for L. bacillatum, L. punctatum (both from Nice) and L. elongatum 
(Falmouth), the type-species of Leptosomatum. 


848 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Some taxonomists have described new species of nematodes in a very unsat- 
isfactory manner. In addition to descriptions and illustrations often being inad- 
equate, the rules and recommendations of the International Code of Zoological 
Nomenclature have not been followed consistently, especially recommendations 
72b, c and d; 73d, 74a, b, c and e, as well as recommendations of Appendix E of 
the Code, i.e., 4, 5 and 19. Indicating paratypes to serve as reference-specimens 
is useful because special attention has been paid to them, but they are most useful 
if deposited in other collections. Care has to be taken not to flatten mounted 
specimens; as stated previously, it is difficult to recognize flattening, and resulting 
artifacts may mislead the observer. 


Acknowledgments 


I am grateful to Mr. P. A. A. Loof under whose guidance this study was con- 
ducted and the L.E.B.-fonds and Wageningenfonds for their financial support. 

Grateful appreciation is expressed to Dr. W. D. Hope for constructive criticism, 
discussion and correcting the text; to Dr. T. A. Platonova for her hospitality, 
valuable discussions and additional measurements; moreover to Dr. W. D. Hope, 
Washington; Dr. J. B. Kirkegaard, Copenhagen; Prof. O. Nybelin, Goteborg; Dr. 
R. Oleréd, Stockholm; Dr. T. A. Platonova, Leningrad; Dr. N. G. Sergeeva, 
Sebastopol; Prof. dr. S. v. d. Spoel, Amsterdam and Dr. R. W. Timm, Dacca for 
the loan of type-specimens. 

I should like to thank Mr. T. S. Ie and Mr. H. Lohuis for advice and assistance 
regarding the E.M.-part. 


Status of Nominal Species 


L. abyssale Allgén, 1951 species inquirenda 

L. acephalatum Chitwood, 1936 probably good species 

L. arcticum Filipjev, 1916 to Leptosomatides 

L. arcticum sensu Mawson, 1958 synonym of L. kerguelense 
L. australe V. Linstow, 1907 species inquirenda; Filipjev, 1918 
L. bacillatum (Eberth, 1863) good species 

L. bathybium Allgén, 1954 species inquirenda 

L. behringicum Filipjev, 1916 species inquirenda 

L. breviceps Platonova, 1967 species inquirenda 

L. caecum Ditlevsen, 1923 to Pseudocella 

L. clavatum, Platonova, 1958 probably good species 

L. crassicutis Platonova, 1958 synonym of L. kerguelense 
L. diversum Platonova, 1978 synonym of L. sachalinense 
L. elongatum Bastian, 1865 synonym of L. bacillatum 
L. elongatum sensu Platonova, 1967 to Leptosomatides 

L. filipjevi Sch. Stekhoven, 1950 synonym of L. bacillatum 
L. gracile Bastian, 1865 synonym of L. bacillatum 
L. gracile sensu Allgén, 1954 to Leptosomatides 

L. grebnickii Filipjev, 1916 to Leptosomatides 

L. groenlandicum Allgén, 1954 species inquirenda 

L. indicum Stewart, 1914 species inquirenda 

L. kerguelense Platonova, 1958 , good species 

L. keiense Micoletzky, 1930 good species 

L. longisetosum Sch. Stekhoven, 1943 synonym of L. punctatum 
L. longissimum (Eberth, 1863) species inquirenda; Filipjev, 1918 
L. micoletzkyi Inglis, 1971 species incertae sedis 

L. pedroense Allgén, 1947 species inquirenda 

L. punctatum (Eberth, 1863) good species 


VOLUME 96, NUMBER 4 849 


Status of Nominal Species (Continued) 


L. ranjhai Timm, 1960 species incertae sedis 

L. sabangense Steiner, 1915 species inquirenda 

L. sabangense sensu Micoletzky, 1930 L. sundaense new name 

L. sabangense sensu Allgén, 1942 synonym of L. bacillatum 

L. sachalinense Platonova, 1978 probably good species 

L. subulatum (Eberth, 1863) species inquirenda; Filipjev, 1918 
L. tetrophthalmum Ssaweljev, 1912 species inquirenda 

L. tetrophthalmum sensu Platonova, 1967 to Leptosomatides 

L. tuapsense Sergeeva, 1973 synonym of L. bacillatum 


Species inquirendae; probably belonging to Leptosomatum: 


L. abyssale Allgén, 1951; Japan, Sagami Sea, 400 m depth. Resembling L. bacillatum. 

L. behringicum Filipjev, 1916; Bering Sea. Resembles L. bacillatum but tail. length equal to anal body 
width. 

L. breviceps Platonova, 1967; Barents Sea. Resembles L. punctatum. 

L. pedroense Allgén, 1947; San Pedro, California. 

L. sabangense Steiner, 1915; Indonesia. Cephalic capsule posterior to papillae. Tail length twice anal 
body width. 

L. sundaense; new name for L. sabangense sensu Micoletzky, 1930; Indonesia. Tail length twice anal 
body width in females, long cephalic capsule (10—12 um) placed anterior to papillae. Short pharynx. 
Ocelli in males far anterior. 


Key to the Valid Nominal Species of Leptosomatum 
1. Cephalic sensilla setiform; cephalic capsule absent in all sexes and stages 


CEA UnGLATUIN=COMPIEX)+: Lek HOE RPL) Ae A ead Bee. 2 
— Cephalic sensilla papilliform; cephalic capsule present in females and ju- 

VEMUES o MAeS PIES Re tie pe No it cae prin ale ne 3 
2. Lens 9-11 wm in diameter; pharynx short; ventromedian pre-cloacal pa- 

PUN ASPKESSMILLOMEAS SMe sd onyerdbn Se Ges pad: Bro ees AY NEAR OE L. keiense 


— Lens 6—7 wm in diameter; ventromedian pre-cloacal papilla present 
5. 00:10 8 8 Giapeeeme eamect lees lr toc ag ale ie Schl gal A LS A L. punctatum 
3. Ocelli at 1 corresponding diameter from anterior end, cephalic capsule 


fOPRESTSUANE “say Lm aa KEP ge ak RO MS Ea se aap L. kerguelense 
— Ocelli at 1.5 corresponding diameter from anterior end; males, if present, 
WHIM UIIECE PH Ali CkCapPSuleuit Arty AU leh ORR unre 8 mS oak 4 
4. Cephalic capsule 10 um; male rare; females big, never less than 10 um in 
feneihen Southemmellennsphenre) 91st eer cen ee ee L. clavatum 
— Cephalic capsule less than 9 wm in length; males not rare; females often 
Sone kere arena OM Tia ch ere Me eae e Teen oe eemercd cn er iNawrat Lee naNn ce en Srl Nene Mmm ee Be A 5 
5. Tail length in female twice anal body width (Indonesia) ...... L. sundaense 


Ww talllenothin adults’ lYsranal body width 4.) es ee ee 
eo ke eae L. bacillatum-complex: L. bacillatum (Europe), L. acephalatum 
(East coast USA) and L. sachalinense (Sakhalin Is.) 


Literature Cited 


Allgén, C. 1942. Die freilebenden Nematoden des Mittelmeeres.—Zodlogische Jahrbiicher (Syste- 
matik) 76:1-102. 
—. 1947. West American marine nematodes (Papers from Dr. Th. Mortensen’s Pacific Expe- 
dition 1914-16, 75).— Videnskabelige Meddelelser fra Dansk Naturhistorisk Forening 1 Kjob- 
enhavn 110:65-219. 
1951. Pacific freeliving marine nematodes (Papers from Dr. Th. Mortensen’s Pacific Ex- 


850 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


pedition 1914-16, 76).— Videnskabelige Meddelelser fra Dansk Naturhistorisk Forening i Kjob- 

enhavn 113:263—411. 

1954. Freeliving marine nematodes from East Greenland and Jan Mayen. The Swedish 
Greenland-Expedition 1899.— Meddelelser om Gronland 107(6):1—44. 

—. 1954a. Uber zwei Tiefsee-Nematoden von der schwedischen Albatross-Expedition 1948.— 

Zoologischer Anzeiger 153:318-321. 

. 1957. Ona small collection of freeliving marine nematodes from Greenland and some other 

arctic regions with reviews and analyses of the compositions of all hitherto known arctic 

nematode faunas. — Meddelelser om Grgnland 159(3):1-42. 

Bastian, H. Ch. 1865. Monograph on the Anguillilidae, or free Nematoids, marine, land, and fresh- 
water; with descriptions of 100 new species.— Transactions of the Linnean Society of London 
25:73-184. 

Baldwin, J. G., and H. Hirschmann. 1973. Fine structure of cephalic sense organs in Meloidogyne 
incognita males.— Journal of Nematology 5:285-302. 

Biitschli, O. 1874. Zur Kenntnis der freilebenden Nematoden, insbesondere der des Kieler Hafens. — 
Abhandlungen der Senckenbergischen Naturforschenden Geschellschaft 9:236-292. 

Chitwood, B. G. 1936. Some marine nematodes of the superfamily Enoploidea.— Transactions of 

the American Microscopical Society 55:208-213. 

1951. North American marine nematodes.— Texas Journal of Science 3:617-672. 
Ditlevsen, Hj. 1923. Sur quelques Nématodes libres (Cétes de Bretagne et Rockall).— Bulletin de la 

Société Zoologique de France 48:178-203. 

Eberth, C. I. 1863. Untersuchungen iiber Nematoden.—Leipzig: W. Engelmann, pp. 1-77. 

Filipjev, I. 1916. Free-living nematodes in the collection of the Zoological Museum of the Imperial 

Academy of Sciences in Petrograd.—Ezhigodnik Zoologicheskago Muzeya 21:59-116. 

1918/1921. Free-living marine nematodes of the Sebastopol area.—Trudy Osoboi Zoolo- 
gischeskoi _Laboratorii 1 Sebastopol’skoi Biologicheskoi Statsii Akademii Nauk (2) 4:1-350 
(1918), 351-614 (1921). [English translation by M. Raveh, Isreal Program for Scientific Trans- 
lations, Jerusalem 1968 (part 1 p. 1-255), 1970 (part 2, p. 1—203).] 

—. 1922. Encore sur les Nématodes libres de la Mer Noire.—Trudy Stavropol’skogo 
Sel’skokhozyaistvennogo Instituta 1:83-184. 

—. 1927. Les Nématodes libres des mers septentrionales appartenant a la famille des Enopli- 
dae.— Archiv fiir Naturgeschichte 91 A(6):1-216. 

Hope, W. D. 1967. Free-living marine nematodes of the genera Pseudocella Filipjev, 1927, Thor- 
acostoma Marion, 1870, and Deontostoma Filipjev, 1916 (Nematoda: Leptosomatidae) from 
the west coast of North America.—Transactions of the American Microscopical Society 86: 
307-334. 

Hope, W. D., and D. G. Murphy. 1969. Syringonomus typicus new genus, new species (Enoplida: 
Leptosomatidae) a marine nematode inhabiting arenaceous tubes. — Proceedings of the Biolog- 
ical Society of Washingtion 82:511—-517. 

Inglis, W. G. 1964. The marine Enoplida (Nematoda): a comparative study of the head.— Bulletin 
of the British Museum (Natural History; Zoology) 11:265-375. 

—. 1971. Marine Enoplida (Nematoda) from Western Australia.—Transactions of the Royal 
Society of South Australia 95:65-78. 

Kreis, H. 1928. Die freilebenden marinen Nematoden der Spitzbergen-Expedition von F. R6mer 
und F. Schaudinn im Jahre 1898.— Mitteilungen aus dem Zoologischen Museum in Berlin 14: 
131-197. 

Linstow, O. v. 1892. Helminthen von Siid-Georgien. Nach der Ausbeute der Deutschen Station von 
1882-1883.—Jahrbuch der Hamburgischen Wissenschaftlichen Anstalten 9(2):1-19. 

—. 1896. Nemathelminthen Hamburger Magalhaensische Sammelreise (Hamburg). 1896, 
22 pp. 

—. 1903. Entozoa des Zoologische Museums der Kaiserlichen Akademie der Wissenschaften 

zu St. Petersburg. I].—Ezhegodnik Zoologischeskago Muzeya 8:265-294. 

1907. Nematoda.— National Antarctic Expedition 1901-1904, National History 3:1—4. 
Loof, P. A. A. 1961. The nematode collection of Dr. J. G, de Man.—Beaufortia 8(93):169-254. 
Lorenzen, S. 1978. Discovery of stretch receptor organs in nematodes—structure, arrangement and 
functional analyses.— Zoologica Scripta 7:175-178. 

1981. Entwurf eines phylogenetischen System der freilebenden Nematoden.— Ver6ffent- 
lichungen des Instituts fiir Meeresforschung im Bremerhaven. Suppl. 7:1—472. 


VOLUME 96, NUMBER 4 851 


Man, J. G. de. 1878. Contribution a la connaissance des Nématodes marins du Golf de Naples. — 
Tijdschrift der Nederlandse Dierkundige Vereeniging 3:88-118. 

1889. Troisiéme note sur les Nématodes libres de la mer du Nord et de la Manche.— 
Mémoires de la Société Zoologique de France 2:182—216. 

. 1889a. Espéces et genres nouveaux de Nématodes libres de la mer du Nord et de la Manche. — 
Mémoires de la Société Zoologique de France 2:1-10. 

1893. Cinquiéme note sur les Nématodes libres de la mer du Nord et de la Manche. — 
Mémoires de la Société Zoologique de France 6:81-125. 

1904. Nématodes libres (Expédit. Antarctique Belge).— Résultats du Voyage S.Y. Belgica: 
1-51. 

Marion, A. F. 1870. Rescherches zoologiques et anatomiques sur des Nématoides non parasites, 
marins.— Annales des Sciences Naturelles 13(14):1—100. 

1870a. Additions aux recherches sur les Nématoides libres du Golf de Marseille. — Annales 
des Sciences Naturelles 14(1):1-16. 

Mawson, P. M. 1958. Free-living nematodes section 3: Enoploidea from subantarctic stations. — 

Report of the British Australian New-Zealand Antarctic Research Expedition (B) 6:407-358. 
Micoletzky, H. 1924. Weitere Beitraége zur Kenntnis freilebender Nematoden aus Suez.—Sitzungs- 
berichte der Akademie der Wissenschaften in Wien (I) 132:225-262. 

1930. Freilebende marine Nematoden von den Sunda-Inseln. I. Enoplidae. (Papers from 
Dr. Th. Mortensen’s Pacific Expedition (1914—16, 53), (edited by H. A. Kreis). — Videnskabelige 
Meddelelser fra Dansk Naturhistorisk Forening 1 Kjobenhavn 87:243-339. 

Newall, D. R. 1976. Volume and ionic regulation in Enoplus communis, Enoplus brevis (Bastian) 
and Heterodera rostochiensis Wollenweber. Ph.D. Thesis: University of Newcastle upon Tyne. 

Platonova, T. A. 1958. Contribution to the nematode fauna of the family Leptosomatidae from 

Kerguelen Island [in Russian].—Informatsionnyi Byulleten’ Sovetskoi Antarkticheskoi Ekspe- 

ditsi1 1955-58, 3:59-61. 

. 1967. Free-living marine nematodes of the family Leptosomatidae from the European arctic 

[in Russian]. — Zoologicheskii Zhurnal 46:828-839. 

1968. Marine free-living nematodes of the family Leptosomatidae from Kerguelen Island 
[in Russian]. Resultaty Biologitscheskich Issledowanij Sowjetskoi Antarkitscheskoi Expeditsii 
1955—1958.—Issledovanija Fauny Morjei 6(14):5—-24. 

1976. New Enoplida (free-living marine nematodes) of the U.S.S.R.-seas [in Russian]. In: 
Akad. Nauk. CCCP, Zool. Inst.—Issledovanija Fauny Morjei 15(23):3-164. 

—. 1978. Marine nematodes of the order Enoplida from the coastal waters of South Sachalina 
[in Russian].— Zoologicheskii Zhurnal (57)4:495-498. 

Schuurmans Stekhoven, J.H. 1943a. Einige neue freilebende marine Nematoden der Fischereigriinde 
vor Alexandrien.— Note dell’ Istituto italo-germanico di biologia marina di Rovigno d’Istria 

2(25):1-15. 

1943b. Freilebende marine Nematoden des Mittelmeeres. IV. Freilebende marine Nema- 
toden der Fischereigriinde bei Alexandrien. — Zoologische Jahrbiicher (Systematik) 76:323-380. 

1950. The free-living marine nemas of the Mediterranean I. The bay of Villefranche.— 
Mémoires de I’Institut Royal des Sciences Naturelles de Belgique (2)37:1—220. 

Seinhorst, J. W. 1959. A rapid method for the transfer of nematodes from fixative to anhydrous 
glycerin. — Nematologica 4:67-69. 

Sergeeva, N. G. 1973. New species of free-living nematodes of the order Enoplida from the Black 
Sea [in Russian].— Zoologicheskii Zhurnal 52(11):1710-1714. 

Ssaweljev, S. 1912. Zur Kenntnis der freilebenden Nematoden des Kolafjords und des Relictensee 
Mogilnoje.— Trudy Imperatorskago St Peterburgskago Obshchestva Estestroispytatelei 43:108— 
126. 

Steiner,G. 1915. Freilebende marine Nematoden von der Kiiste Sumatras. — Zoologische Jahrbiicher 

(Systematik) 38:222-244. 

1916. Freilebende Nematoden aus der Barentssee.— Zoologische Jahrbiicher (Systematik) 
39:511-676. 

Stewart, F. H. 1914. Report on a collection of freeliving nematodes from the Chilka Lake on the 
East Coast of India.— Record of the Indian Museum 10:245-254. 

Stiles, C. W., and A. Hassal. 1905. The determination of generic types and a list of roundworm 
genera, with their original and type species. — Bulletin of the Bureau of Animal Industry United 
States Department of Agriculture 79:1—150. 


852 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Timm, R. W. 1953. Observations on the morphology and histological anatomy of a marine nematode, 
Leptosomatum acephalatum Chitwood, 1936, new combination (Enoplidae: Leptosomatin- 
ae).—American Midland Naturalist 49:229-248. 
1960. A new species of Leptosomatum (Nematoda) from the Arabian Sea.—Journal of 
Helminthology 34:217—220. 
Villot, A. 1875. Recherches sur les Helminthes libres ou parasites des c6tes de la Bretagne. — Archives 
de Zoologie Expérimentale et Générale (1) (4):451—482. 


Department of Nematology, Agricultural University, P.O.B. 8123; 6700 ES, 
Wageningen, The Netherlands. 


Appendix.— Numerical data new records. 


Abbreviations: Sex (S) or stage; specimen number (SN); length (L); distance to lens (DL); distance 
to nerve ring (DNR); pharynx length (PL); caudal length (CL); cephalic width (CW); width at level of 
ocelli (OW); at nerve ring level (NW); pharynx base (PW); vulva or midbody width if not applicable 
(MW) and anal body width (AW). 


S SN L DL DNR PL CL CW OW NW PW MW AW a b c Vv 


L. bacillatum, pop. 1; Den Helder 


1072-bY 2960" 48) 1829 730)" “6b E7228) 35" 41 390 28) 2 aa 
1063-b© “6760 92-303" 1214" 77-31) 58) 75° 83" 86953-8195 E ES Geo omoU 
1064=a 7250) "803 UP A214) 80930 559970 83) 889] 55" Sey EO: Ono nae 
1065 FESO" 86 "297 126” “9629 52 6467 O66 853 hse Gea 
HOP3=D TIZOW 849283 1118" 683293153" 72" 9836 8S) 66) FCO One 
LOW3-CY = 859098" 88296" W666" 86931 S89 742° 7 83183 59M 103 aeaenew 
1062 $690 °° 84-0324 PTS7. 99 30" S274" 2784 86564" “10 eae 
1070 S760 388 3271345586 349 66789110 1359 6609 65 Osmo Za 
O74 103230) 102" 333) V476" 19199360166" “S6) 106127) 68" —SieG ome aso 
1069 10,870 108 342 1378 94 34 64 91 114 128 67 85 7.9 116 57 


. bacillatum, pop. 2; Kattendijke 


LOS2-ay, 2470) S0..199) (669) 4658. 17.30: 36, 384.37... (30) 4 65 Ses 
1LO82-¢ 2730) 55, 172, S567. 64 16) 31.41, 47 4N 28 “S8iy aoe 
NOS4-ay 3140) 61) 200) 3709 9635 922-41) 52) 56) 52>, 39 Ge a) 
1054 5340), Sly 292) TNS 974, 252045, 59) (64... 164). 47. 83 7ee ae See 
1055 J300 \ See 69 ED ee See 28e50) ls ASO 5 SG.) 9) lee GsCmmme 
1052 7540) T3927 VASA 81355, 77 28). Sil. S69 Se G ee meet 
1056 81309 83.283, 1345388433. 59.. 72) Sly 83.) 55 998s sooo 
1045 9080), 89° 245, 1287, 84,31 53, 70) e738) 980) 59 ia eles 
1050 9120.78 286 1378 88 31. 53.74 389) 89 58 1025.66 eeI0 tae? 
1049 105370\ 75° 306 1378 9:86.28. 52 267% 78.) 718.056. «133 seu 


. bacillatum, pop. 3; Burghsluis 


2152 6270) "82" 288" 1123 "S33" 68 975 Sil" Sl 58" 77 = Ome 
2165 1040” 89 283 1251" 8934645 S75 100) 96" 6259/0 Crema 
2037 T3590" 90) 333) SV192" "9393 S56" Dee 83") ONIN OS 59) le On Smmmoe 
2162 1570" 78 331 "1281 815933 S85 718 86 Sil Ol 88m =o emoommas 
2008 T3810" “S2" 350 WS8) Ol 30 54" 0 iS eG) ON N04 eGo eis 
2047 SOSOm 82) 321 IDS8) Y87e33" 253) | e751 9) 2 ae ee Oe On mem 
2026 $330) 78 327 122589" S30 SG) moles FSO lente oleae Ome 
2077 9060" “7953457 TA 93-30" Sie 65) 73) 1S) Od Oreo vam 
2018 O1GOe “S77 3Tl 1254" 98437 72 999" TOO 110) 645" 83 aes Ooms 
2064 10/440 101 322° 1336 "98" 34 "64" "90 L0G tis 72S es) HO Memes 


munzezzenznrnh ZnZznnZzeuuw4 aan nrzezeen74 


VOLUME 96, NUMBER 4 


Appendix. — Continued. 


853 


Abbreviations: Sex (S) or stage; specimen number (SN); length (L); distance to lens (DL); distance 
to nerve ring (DNR); pharynx length (PL); caudal length (CL); cephalic width (CW); width at level of 
ocelli (OW); at nerve ring level (NW); pharynx base (PW); vulva or midbody width if not applicable 
(MW) and anal body width (AW). 


Cw OW NW 


S SN it DL DNR PL CL 
L. bacillatum, pop. 4; Texel, ‘t Horntje 

M 3108 6430 84 316 1242 89 30 
F 3074 7480 74 286 1112 86 34 
F 3158 7790 78 297 1177 78 34 
M 3106 8040 88 302 1057 81 31 
F 3024 8330 94 347 1273 91 35 
M 3056 8750 89 310 1196 86 32 
ESO 30). 925 3141335) 78) Sil 
F 3046 11,780 92 330 1483 101 34 
F 3053 12,820 91 337 1464 83 35 
F 3014 13,570 91 326 1446 82 34 
L. bacillatum, pop. 5; Texel, Oudeschild 

J 1012-c 1370 47 149 407 52 13 
J Oily Bas V4 BID Wiz) 2 27 
J 1061 4260 63 227 896 66 23 
J 1003-a 5330 70 247 952 74 28 
J 1010-b 6690 78 277 1062 66 27 
M 1004 7080 90 288 1155 81 31 
M 1006 7770 »=6°89 6312 1195 80 33 
M 1035 8040 86 335 1378 86 31 
M 1033 8920 95 336 1336 78 32 
F 1009 9350 78 274 1148 78 33 
L. bacillatum, pop. 6; Wimereux 

F A-59 11,560 89 279 1420 83 27 
L. bacillatum, pop. 7; Ambleteuse 

F 1277-1 7980 83 284 1048 79 25 
F 1277-2 8570 109 329 1284 77 30 
L. bacillatum, pop. 8; Banyuls 

F 1286-1 7620 102 288 1155 89 33 
F 1286-2 13,490 109 376 1741 89 35 
F 1286-3 16,890 135 415 1892 106 33 
L. bacillatum, pop. 9; N.E. England 

M 76101 8720 82 324 1342 93 29 
F 76102 10,190 91 294 1305 108 32 
L. acephalatum, pop. 1; Mexico, N.E. Asc. Bay 
J 76103 4700 69 227 779 76 25 
F 76104 7200 95 272 997 133 38 
J 76105 7290 101 300 1070 91 31 
F 76113 7780 97 303 1088 95 32 
F 76106 7790 108 220 1178 90 37 
M 76107 8030 75 337 1132 98 35 
M 76108 8430 101 360 1142 92 32 
F 76109 8970 112 316 1124 96 34 
L. acephalatum, pop. 2; Mexico, Allen Point 

F 76110 7720 82 298 1106 103 37 


PW 


112 


122 
121 


130 


MW AW 
95 66 
109 66 
33° 181 
86 =665 
Di OF 
a 0 
Bz. WO 
WZ WZ 
146 70 
136 8673 
24 22 
We eal 
53 3 
81 47 
99 62 
D2 19) 
92 66 
84 664 
105 63 
107 64 
156 671 
114 64 
KO @il 
134 73 
94 «264 
I) OF 
oo OY) 
113 66 
1B 52 
107 68 
112 81 
115 676 
110 69 
118 73 
98 66 
140 77 
IW 72 


8.1 


7.6 
6.7 


6.6 
Tol 
8.9 


6.5 
7.8 


6.0 
Tes) 
6.8 
V2 
6.8 
7.1 
7.4 
8.0 


7.0 


101 


114 
120 


139 


58 


60 
59 


60 
59 
61 


59 


58 


57 
60 


57 


56 


854 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


Appendix.— Continued. 


Abbreviations: Sex (S) or stage; specimen number (SN); length (L); distance to lens (DL); distance 
to nerve ring (DNR); pharynx length (PL); caudal length (CL); cephalic width (CW); width at level of 
ocelli (OW); at nerve ring level (NW); pharynx base (PW); vulva or midbody width if not applicable 
(MW) and anal body width (AW). 


S SN L DL DNR PL CL CW OW NW PW MW AW a b c Vv 


L. acephalatum, pop. 3; Mexico, N.E. Asc. Bay 

76092 4660' 86 188 852 77 29) 54 73 79 77 52) 59°55) ol 
76093 7630 106 326 1051 90 32 60 77 85 92 60 83 7.3 85 
76094 7790 102 333 1178 89 37 70 103 130 153 77 Sil 66 88 56 
76095 7810 110 343 924 92 34 62 81 100 99 67 78 85 85 
76096 8090 118 341 1106 109 34 70 98 123 123 74 66 7.3 73 50 
76097 8700 87 358 1250 96 32 65 90 113 139 74 63 7.0 91 58 
76091 9860 110 343 1215 91 30 62 96 126 147 81 67 48.1 108 54 


L. acephalatum, pop. 4; Mexico, Niccehabin Reef 

J 76098 1220) 95 DS DIO B85 We 52 Sl M07 120 Wl Ol 7.5 8 
L. acephalatum, pop. 5; Mexico, Suliman Pt. 

J 76099 S430) 65 . 322 562 10 2 a) 65 BF DS SGC‘ SLID 


L. acephalatum, pop. 6; Mexico, S.E. Cozumel Is. 
EF 76100) 10170 “878 316 Tis T0840) 79) 117 164" 19355 Oe S53 SkG eo aa 


9] lal "ol SS ol SS 


L. clavatum, Macquarie Islands 


76078 GOS0 MOS B37 NS MOG 25 CS si Sys 8) 3s 8 oA ST 
76079 6520) 85) 27ly NOMA 19425) 58) 81) (85) 10255 67 SOTO sto? 
76080 YOO WG 370 M52 12S 37 C2 INO IWS Wee Sy S27 6.2 66 
76081 SS7ON Sil SIO OA. WD 27s 52) 3 a Sie Oe eS ee 
76082) 11,050) 132, 427 1386125 36 8) 106 119) 133) 89) 83) 8:0 esse” 
76083 12,120 113 374 1589 121 31 82 89 102 114 74 106 7.6 100 63 
76084 12,510 133 452 1673 100 33 83 110 125 144 88 87 7.5 125 62 
76085 12,780 141 419 1606 142 38 85 110 127 154 92 83 80 90 63 
76086 13,030 146 444 1741 146 40 85 108 125 158 94 82 7.5 89 63 
IOS) IBASO Wl 43 1639 152 B35 7 ys ie Mit OS) WD 3.2 se ao 
76088 | 13,960) 121 374 758 M42 33°83) NO 134° 162) 93 86) ee o Seon 
76089 14,230 150 469 1656 139 37 85 114 150 168 92 85 86 102 61 


ecMiesMics Mie: Mesias Ie siies i= i 


L. keiense, pop. 1; Philippines, Telbang Cove 

J 76039 3540» 65) 1196 544 77 19 46, “61 72) 79 52). “45 esse 

F 76040 ASTON 84) 256) 8384 ~ 90) 24,7445 7 8 95) 58) 2 Ono 
M 76041 5780 7 203) 3689 90-3253) 7 T4602 Soe or 


L. keiense, pop. 2; Philippines, Cangaluyan Is. 


J 76042 1710 42) 148) 406 46) 6, 32) 44 47) 48 8 eso 
J 76043 ASS Ol ue 29 592 Sle 59) Sin Soe Or en eee) 
M 76044 G80) *80 9273) 834" 66 25° 445s 8h 645" CI 55) 97 eee 
M 76045 6580 68 278 888 63 27 45 60 60 63 55 104 7.4 104 
F 76046 7870 67 280 979 69 25 48 71 90 114 54 69 8.0 114 
L. keiense, pop. 3; Philippines, Bolic Is. 


J 76047 4830 76 222 558 82 27 AQ <69) 77.85.) 500) 57 aoe ms, 
J 76048 SOO) 92256, 1642) 98 27. C489 OS 2 le CoS eer) 
M 76049 120 89 BO 689 NOS 37 G2 Bil 88 93 © G2 WO 7 


L. keiense, pop. 4; Philippines, Gr. Santa Cruz Is. 
J 76050 AO) SS QM 503 6 22 450 GHOST) OO DDO 


VOLUME 96, NUMBER 4 


Appendix. — Continued. 


855 


Abbreviations: Sex (S) or stage; specimen number (SN); length (L); distance to lens (DL); distance 
to nerve ring (DNR); pharynx length (PL); caudal length (CL); cephalic width (CW); width at level of 
ocelli (OW); at nerve ring level (NW); pharynx base (PW); vulva or midbody width if not applicable 
(MW) and anal body width (AW). 


S SN L DL DNR PL CL 
L. keiense, pop. 5; Philippines, Sacol Is. 

J 76051 3770 §©663 182 490 75 
L. kerguelense, pop. 1; South Georgia 

M 76052 7150 64 452 1031 £87 
J 76053 7420 69 386 1166 73 
M 76054 8530 68 345 1169 100 
F 76055 9350 67 408 1268 125 
M 76056 10,210 77 431 1082 106 
F 76057 15,840 73 469 2315 116 
L. kerguelense, pop. 2; Macquarie Islands 

J 76058 10,710 64 366 1470 107 
L. punctatum, pop. 1; Red Sea 

F 76037 3660 67 213 816 81 
L. punctatum, pop. 2; Banyuls 

J 1286-4 2940 58 214 608 69 
F 1286-5 7600 79 321 1065 114 
M 1286-6 7830 85 319 991 106 
L. sabangense; Red Sea 

J 76038 6490 58 213 761 95 
L. sp. A; Macquarie Islands 

J 76067 7610 71 267 879 98 
J 76068 8720 70 275 896 95 
J 76069 8790 74 292 930 96 
F 76070 11,100 71 284 1048 115 
F 76071 11,830 75 366 1166 116 
F 76072 12,150 86 325 1031 120 
F 518 12,290 81 370 1200 116 
F 504 12,440 87 370 1268 114 
F 76073 14,090 75 362 1301 129 
Syringonomus typicus;, Recife, Brazil 

J 76059 3310 — 239 667 #4=279 
J 76060 3330 — 241 666 £65 
F 76061 4450 — 280 881 101 
F 76062 5050 — 321 952 104 
F 76063 5240 — 313 904 118 
J 76064 5360 — 297 762 95 
J 76065 5570 — 305 928 110 
F 76066 6880 — 453 1227 110 


CW OW 


50 


NW 


69 


102 
129 
112 
116 
112 
14] 


112 


58 


57 
87 
99 


48 


67 
7\ 
67 
WY 
108 
85 
79 
73 
76 


56 
61 
70 
71 
74 
71 
63 
66 


PW 


78 


108 
152 
125 
160 
124 
162 


146 


66 


69 
99 
105 


52 


VI 
81 
V7 
102 
102 
112 
94 
85 
94 


66 
73 
81 
88 
89 
82 
74 
84 


MW 


84 


116 
156 
145 
168 
134 
201 


162 


63 


61 
111 
114 


61 


92 
99 
83 
139 
112 
139 
133 
110 
108 


78 
81 
95 
113 
116 
95 
89 
100 


AW 


59 


79 
114 
91 
119 
92 
135 


106 


106 


83 
88 
106 
80 
106 
87 
92 
113 
130 


42 
41 
47 
45 
45 
56 
63 
69 


5.9 
6.4 
V3 
7.4 
9.4 
6.8 


7.3 
4.5 


4.8 
Tak 
7.9 


8.5 


8.7 
7 
9.5 
10.6 
10.1 
11.8 
10.2 
9.8 
10.8 


5.0 
5.0 
5.1 
5.3) 
5.8 
7.0 
6.0 
5.6 


102 


WS. 36 


37 oY 


100 


45 54 


43 
C6 
74 


68 


856 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 


BIOLOGICAL SOCIETY OF WASHINGTON 
PROCEEDINGS 


110th Annual Meeting, 10 May 1983 


The meeting was called to order at 2:35 PM by President Paul Spangler. It was 
moved and passed to dispense with the reading of the minutes of the previous 
meeting. 

The Treasurer’s Report was offered by David Pawson for Treasurer Leslie 
Knapp. Pawson noted that the Society’s net worth is now increasing because of 
the increased page charges and the reduction in the number of free pages. The 
discrepancies between the 1981 and 1982 items on the balance sheets are attrib- 
utable mainly to calendar differences, i.e., timing of payments and receipt of 
income. A motion to approve the Report was passed unanimously. 

The Editor’s Report by Editor Brian Kensley was offered by Acting Editor 
Stephen Cairns. Cairns added that we have received 43 MSS so far in 1983. 
Volume 96(2) is in galley and 96(3) will be sent to Allen Press shortly. Ten MSS 
have been scheduled for 96(4). A motion to approve the report was passed unan- 
imously. 

The President announced the results of the annual elections: President, David 
Pawson; Vice-president, Donald Davis; Treasurer, Leslie Knapp; Secretary, Cathy 
Kerby; Councilors Daryl Domning, Carl Ernst, C. W. Hart, Robert P. Higgins, 
David A. Nickle. He then turned the chair over to incoming-president Pawson, 
who led a round of applause in appreciation of the services of outgoing-president 
Spangler. 

The meeting was adjourned at 3:01 PM. 


Thomas E. Bowman, 
Substitute Secretary 


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CONTENTS 


Three new species of Brueelia (Mallophaga: Philopteridae) from the Mimidae (Aves: 

Passeriformes) N. Sandra Williams 
Description and phylogeny of /saacsicalanus paucisetus, n. gen., n. sp., (Copepoda: Calanoida: 

Spinocalanidae) from an east Pacific hydrothermal vent site (21°N) A. Fleminger 
Eight new species of Indo-Pacific crabs from the collections of the Smithsonian Institution 

Tune Sakai 

A new species of serranid fish genus Plectranthias (Pisces: Perciformes) from the southeastern 

Pacific Ocean, with comments on the genus Ellerkeldia 

Phillip C. Heemstra and William D. Anderson, Jr. 
Redescription of the Brazilian labrisomid fish Starksia brasiliensis 
J. T. Williams and A. M. Smart 

Description of a new species of Echiodon (Teleostei: Carapidae) from Antarctic and adjacent 

seas Douglas F. Markle, Jeffery T. Williams, and John E. Olney 
Teleostean otoliths from the late Cretaceous (Maestrichtian age) Severn formation of 

Marvland Richard W. Huddleston and Kurt M. Savoie 
A new subspecies of fox sparrow from Alaska J. Dan Webster 
A revision of the Golfingia subgenera Golfingiella, Stephen, 1964, and Siphonoides, Murina, 

1967 (Sipuncula) Edward B. Cutler, Norma J. Cutler, and Peter E. Gibbs 
Two species of 7ylos Audouin from Chile, with notes on species of 7y/os with three flagellar 

articles (Isopoda: Oniscoidea: Tylidae) George A. Schultz 
Bonaducecytheridae McKenzie, 1977: A subjective synonym of Psammocytheridae Klie, 1938 

(Ostracoda: Podocopida: Cytheracea) K. G. McKenzie 
Heteropyramis alcala and Thalassophyes ferrarii, new species of Clausophyidae (Calycophorae: 

Siphonophorae) from the South Pacific Angeles Alvarifio and Kenneth R. Frankwick 
Caecidotea filicispeluncae, a new troglobitic asellid isopod from Ohio 

Thomas E. Bowman, III, and H. H. Hobbs, III 
Pycnogonida of the western Pacific islands II. Guam and the Palau islands C. Allan Child 
Recognition of two species of double-lined mackerels (Grammatorcynus: Scombridae) 
Bruce B. Collette 

Two new species of coral toadfishes, family Batrachoididae, genus Sanopus, from Yucatan, 

Mexico, and Belize Bruce B. Collette 
A revision of the Seguenziacea Verrill, 1884 (Gastropoda: Prosobranchia). I. Summary and 

evaluation of the superfamily James F. Quinn, Jr. 
Observations on species of the fossil genus Axopora (Coelenterata: Hydrozoa) and its evolutionary 

significance to the Stylasteridae Stephen D. Cairns 
On a small collection of entocytherid ostracods with the descriptions of three new species 

Horton H. Hobbs, Jr. and Auden C. McClure 
Seven new species of the Indo-Pacific genus Eviota (Pisces: Gobiidae) 
Susan L. Jewett and Emest A. Lachner 
Revision of the genus Leptosomatum Bastian, 1865 (Nematoda: Leptosomatidae) 
; Tom Bongers 


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