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Ee oc a AAS a Vi, J aa 2 - > =. 7 hee m AS g m eld aa B RARI ES SMITHSONIAN INSTITUTION NOILALILSNI _ NYINOSHLNS a7 iuyy i = at = = = 4 et re 4 je ~ 4, NX e : ae a O ED 2) =. ee baie 2 om ip \ = i 2, at ae ee eS > >’ ( Ne Be er ge a any JILNLILSNI NVINOSHLINS S3I1YVYdiIt LIBRARIES SMITHSONIAN INSTIT! sate c 2 i Pa ” Ww o tl na a cx aa oe 5, ey o _< vad Ea re * Ke = ad te ”) pan 7p) an w = w = w = OILNLILSNI . x Ns z z < = < Z “4 z = FA re) = aa ae a oH : B 2 é 2 = 2 E 2 = = > = > = SMITHSONIAN INSTITUTION NOILNLILSNI NVINOSHLINS S3IU‘ _ S58 a Pee ” _ nw S XA tok «. i fg a = AS 2 = “Gite 5 : eB = m “ip = 28) = < m me " > e z iY ad OILALILSNI NVINOSHLIWS 54 1yYVvuad ie LIBRAR! ESi SMITHSONIAN NSTI | Te aes on ial re CQ Seen oe Pe enn ioy vont ees, ee eee INSTITUT saluva INSTITUT INSTITUT NLILSNI NVINOSHLINS SAIYVYE!II_LIBRARIES SMITHSONIAN RARIES SMITHSONIAN INSTITUTION NOILNLILSNI NVINOSHLIWS NVINOSHLINS S3IYVa' SMITHSONIAN NVINOSHLIWS DQG" LY SMITHSONIAN S LIBRARIES xs S3l1u¥vVusiT LIBRARIES SMITHSONIAN _INSTITUT! INSTITUTION NOILNLILSNI SJIUVUGIT LIBRARIES INSTITUTION NOILNLILSNI S3IYVUGITLIBRARIE S31uvyus!l) ZARIES SMITHSONIAN INSTITUTION NOILMLILSNI_NVINOSHLINS S3IYVUE ” z w” z w r = = = = = YY =f x we: 5 oN = Ae 8 OA re) =z 2X Ws : 0 LF &. 2 E NY" 2 = 2/7 = = ra p>" = > a ” ei = ” i _NVINOSHLINS S34 IYVUYUGIT LIBRARIES SMITHSONIAN INSTITUTI = = ne Zz 7 ud ul g % z - Ye” =f ae 4 “fo A es = = < EE igo A, er ~ Yar om — mom.” = Su 2 = S RAR I ee NS KITUTION NOILNLILSNI NWINOSHLINS S31YVu! = ox ic re) re S) = Ne fe) i tes E pe = be as 2) : WS = Ke : > tail Ey = E os a eS OB es o = D . = fs a) = w _ & Peon SVNRESNS 4 1uVvuag riot po MITHSONIAN Feduccia, Alan. Comments on the phylogeny of perching birds ___-__----_---_-_______- Flint, Oliver S., Jr. Studies of Neotropical caddisflies XXIII: new genera from LE SNES Cy dees ee) se Ses de a A ee A Se es Flint, Oliver S., Jr., J. Reese Voshell, Jr., and Charles R. Parker. The Hydro- psyche scalaris group in Virginia, with the description of two new species (Trichop- REBT AROTI SNC LO AC) Meee im te ht Andis Veena Rol vane ye oe he PI ¥ soe ee Nahe ao) Pehle Goy, Joseph W., and Anthony J. Provenzano, Jr. Juvenile morphology of the rare burrowing mud shrimp Naushonia crangonoides Kingsley, with a review of the genus Naushonia (Decapoda: Thalassinidea: Laomediidae) _________________- Haman, Drew. Voorthuyseniella ventressi, new species (Microproblematica) from mea SaieletstOcene Or I ouisiana- smi ooh 100 rat IY Lone ae sO ge Heaney, Lawrence R. A new species of tree squirrel (Sundasciurus) from Pala- wan Island, Philippines (Mammalia: Sciuridae): 0) ee Hendler, Gordon. Sex-reversal and viviparity in Ophiolepis kieri, n. sp., with notes on viviparous brittlestars from the Caribbean (Echinodermata: Ophiuroi- Hobbs, Horton H., Jr. A new-crayfish from the Ouachita River basin in Ar- Rane ccapodanCambaridae), 222. aan Tete? YE on ta Poe iets oye h Hobbs, H. H., III. Additional notes on cave shrimps (Crustacea: Atyidae and Ealaemonidae) from the Yucatan Peninsula, Mexico .__.....--.-..-_.----- Humes, Arthur G., and Masahiro Dojiri. Poecilostome copepods (Lichomolgidae) from the alcyonacean coral Cespitularia multipinnata in the Moluccas ____________ Karaman, Gordan S. Two new species of the genus Jdunella Sars, 1895 (Crus- tacea: Amphipoda) with remarks on the other species. (Contribution to the mmenecdse ofthe Amphipodaro4) es. oru) i Oeeee 3. 4 ewe fae 8 eb Karaman, Gordan S., and J. Laurens Barnard. Classificatory revisions in gam- MeMecanoAmphipoda (@nuistaced). Part dint see tees wane yee ees Karaman, Gordan S. Stenocorophium bowmani, a new genus and species of the family Corophiidae from the Palau Islands (Crustacea: Amphipoda) ________________ Kenk, Roman. Freshwater triclads (Turbellaria) of North America. XI. Phago- cata holleri, new species, from a cave in North Carolina _______________-____________ Kensley, Brian. Redescription of Haliophasma barnardi (Monod) from West mance (© rustacea-)lsopoda: Anthuridae) ore: 2). eifion) 6) een ae Kensley, Brian. Redescription of Zonophryxus trilobus Richardson, with notes on the male and developmental stages (Crustacea: Isopoda: Dajidae) ______________ Kensley, Brian. New species of Anthurideans from the Cook and Fiji islands eeanstacca-lsopoday Anthuridea) swe Our heen amends Wien tap Sepia Ul Ls 132 Cerapus:benthophilusin seas ele Oe i Boa EN sek ee eee 98 Ctenocheles:leviceps: s220-t.000 0s eee. oe) 0 a cel See ee 295 Cypsiphimedia ‘stegosaura:iacs 24. {eo ee Ee eee 108 Dactylopleustes 2 221 sheer a oe i eV Le ies ee ee a ee 112 EObroleiisoe 22 es ee gee) er ane Gee Toa * Ae 376 OVO) Gh, eae eee ae one ane TEE ie a OE AD MBIT AM ee ee 375 Foxiphalis 2a ote Bt NN EES ae RUS ie es 372 Gintphianeus < ky) vec eee ae) oo SL a ee ee 150 Gonodactylus albicinetus: 22.50... 2. Bo Oe ee ee 634 GOSTGTICENSIS: SS... 2.2 es ote ee Se 636 (GiOTIGUOXUS 28 od ose oN et eer s oe oe ee Maes ae 374 IdunellarbowenGe 2.254 «0 ec We oe a ee ee ee i ee qs MOQ OUUPE IR Stee nf ee eal ae ee ee ees ale el 81 ISI ELING ec MO rom sot Ve lh Se Ca a ae Se 133 WOWVECOSOIMINGTUS: <2) et ee ee 134 MMesanthura GHildi <2 e 817 Mesosignum antarcticum ot ee ee See een ee eh 328 IMIIGIIDIUS 22 2)! a ee) ee 400 (2) i ne reer RMS ROM Nee wete Mew Seek 400 INannosquilla canica 2.020). a 637 VOSQUCZL... 2 sie oes eee ee 380 Matalanthura fijiensis. 8 2 en fo a ee ee 820 IPAMUNYIRAVCOUGHIS -cco% coco ene ee 823 PAMAMLOIL WS CXCCH GENS aeccses cre seo a ee 53 QUA CGTIS 5h ooo a osteo isa AS aA ce tn see 56 Paranthura.astrolabium —_.....-..--<2--.- eee Se 830 WOWNESICO 2.235. oo ae eee eee 833 banapleustes COMIger N.C. 2s s-- ee ee 113 APATAG N.C. a2 ee 113 Parapleustes? euacanthoides n.c. MRPPSIAICITLCTIIES) = Seine Ret 9k Se ee ee ee Ee Ne 254 ER RETECUIOICET VLC es po Rit Wee a ge Al ee FS a 256 an CCRP ISIS MIC INV tees ere ee eh eee Pe EA Be alway we ok eRe 209 em ANTERIERR AU CLE OLONL Giese ee a oe ee Ve EEE Ds | ee a iy ed 827 SNIREAOUETOR FAMILIES Petites ciatese edu Sate eee hk Se a ek ee ed) as 769 NS IES DLT A EIEN SIRENS SIE NUE Die ie, SEAN I een Neate 2. a oe 769 Seen eeeNts DRUIS CLIICL fy ete et he eee Re pe ah ne ee ee 804 PIECE SOD Sooo eee Sneek See re ee ee OE ee a 151 ER TERT ASSES TOR AOOTASIECL CAD seat ears SR Sar ain ca Sah Ses © rh? at ee 154 * PED CSRNCUIIGN Se ce SON a Jee ai Ae Sane eae an re a Bens a ee See ees eee 371 EINE cia ent end a S| Tops LI Be fee re me 141 PEUGEOT ET ei a a Aa A a NN On iS 2 a es PC 184 EERE DUTT eR IS AT APN SITE 2 3 0S a cl 580 PDD ea TP Tac es Vale ae Sree enn el NR a ae LS Ber 581 _ ARES CARGO TS Ses NEE Se IO GTS ae i eee ae ee eee eet eee eer. 143 NEMEC CHEN ECS ese se es re ee ORS AN ea RM Ol oe ha es a a ee 113 ws PEE CET os SE INE ER Cr EE ORS RF Oe a PORE PO Meee OR ee 137 (UEDA DID IN ae eis Seapets Ne gt A ai EIT NG ia A eS OP SR A Eh AA a RO NN 149 oD PAQUET) ese Seca Bee Mir sg A ee I A Aa a 138 DETRPGLE Le Bee ee ere, A es MEU EY Sg et 139 POE CHAC CIS 2 22k Be 28 SP dn Sn a 64 Diplopoda Delophon holti _______-_--____-- ae fe ee AME EN ee Sg ANS Ss UR Sh 544 Insecta EMME EIDE CTL OTE CS al x RMR Sek LEO mth Bi EIA Ar PE A hy BS cn saat ete 646 BEDE ECT TTI ccs BE ING OS CASE RE RTS Te yl SMe | Der Waee Neha ea en epee ae Te Meee 649 DELGTIIS atSaoae 2 Ngee I SIN LE ent Hl BN, ee Ref oO RRR GE gO OR AE ce 646 LEP DUEL ORE IVT alae a 0 a ON OSI gee J erie” See of se eR oS rs 643 RGAEE CUTS Cusp ce Sok i hee eS tA Oe eG Oe Oe ee a Ma oe 644 IRE EN Re AEE cE ell ee ee EES ee een me Lee eee eetlas 698 [LIA OE Se Bh a A ue AE RE A Ae EAL ait RR, VS cen Peter te en he 700 DOP DS LS SORE ee ne TES Die oP Ald ee Wf eet eg ea eR 333 on CED ERIAOIS G Ss e o 0 a eR en ee NTI, 28 Sale a ke SR bE a EO 753 GECTAMO VELEN Ieee SOE ON OR RO eo eee tl nen be Dib 754 PERRIS ELIS OOLECEC IAS LLE Ss Nec tema ei hs Lae see ME, Re Ne IN ie eh ee 289 STE T IT ATTA POS OI NES OUST sy A dL a a Prk ese eR ve eet 287 EMERG SN CUGCIDIS ST mete oa Sat ik eo Kay bh se Ue ee aro a 842 ADICTS | oe Eg RD Seas | MEL Tee ene SE geo an es We BOE eee IES OR erin eae ee Ae 854 DECRIED The eet AE AE IR SS SE Ta oe sep SAE LEO et gE I aa aeRO re 641 RP CTICHIC KIC IIS Stee Sate eae « Rie eater AS te een OS oe Nena, he ae 641 ECHINODERMATA Asteroidea UREA CSIAS UG aL ern ea eee nh nies Mes SY Leesa Pes SP ee Pee EE es eb ES 8 oN ee 801 EAUPRAN ASEC TSP CHICIES. (eee naten ieee 1 Shek a Oe eee es ee oe 70 Echinoidea Wlypeaster Kieri -.... SSN APs Wace ORG ey EO ee NOCH Ne Wy Wit es Se See Bs Re a 796 xi Ophiuroidea Ophiolepis Kieri 00 2--2 28 ee ee 784 CHORDATA Pisces GOSINOCGINDUS 2225) 22562 ae ee 2 674 INOLIOCGINPUS ive 8 ice 280 ee es 482 Synenathus.caribbaeus =. 222222252 bk ee 672 Swaodus doaki 2.22.250-0. 2! 2) ek pe ee es 166 JOCUMUI: | a8 oP 2 a Ee MD eo Le 169 rubromarmoratus 2.22 2 ot A ee eee 172 Amphibia Pleutherodactylus loustes’!)! 02.00 8 on ee ee 498 Reptilia Diploglossus anelpisius 20. ...2.2.1ee2 io) 1 ee ee 3 eiocephalus rhutidiya 222202. noo ee ee ee 273 Mammalia Sundasciumis' 7abori. 2222200010. ee ee ee eee ee 281 Xi Proceedings : of the _ BIOLOGICAL SOCIETY - - WASHINGTON Volume 92 18 July 1979 Number 1 THE BIOLOGICAL SOCIETY OF WASHINGTON 1978-1979 Officers President: Clyde F. E. Roper Secretary: Michael A. Bogan Vice President: Oliver S. Flint, Jr. Treasurer: David L. Pawson Elected Council Richard Banks Robert H. Gibbs, Jr. Arthur H. Clarke William R. Heyer Brian Robbins PROCEEDINGS Editor: C. W. Hart, Jr. Associate Editors Classical Languages: George C. Steyskal Invertebrates: Thomas E. Bowman Plants: David B. Lellinger Vertebrates: Richard Banks Insects: Robert D. Gordon Membership in the Society is open to anyone who wishes to join. There are no prerequisites. Annual dues of $7.00 include subscription to the Proceedings of the Biological Society of Washington. Correspondence concerning membership should be addressed to the Treasurer, Biological Society of Washington, National Museum of Natural History, Smithsonian Insti- tution, Washington, D.C. 20560. The Proceedings of the Biological Society of Washington is issued quarterly in February, May, August and November. Manuscripts, corrected proofs, editorial questions should be sent to the Editor, Biological Society of Washington, National Museum of Natural History, Smithsonian Institution, Wash- ington, D.C. 20560. Known office of publication: National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560. Printed for the Society by Allen Press, Inc., Lawrence, Kansas 66044 Second class postage paid at Washington, D.C., and additional mailing office. PROC. BIOL. SOC. WASH. 92(1), 1979, pp. 1-9 A NEW SPECIES OF DIPLOGLOSSUS (SAURIA: ANGUIDAE) FROM HISPANIOLA Albert Schwartz, Eugene D. Graham, Jr., and Julian J. Duval Abstract .—Diploglossus anelpistus, a new species of anguid lizard from the Antillean island of Hispaniola, is most closely related to D. warreni, but the known distributions of the two species are separated by a distance of 300 km. The two species do not differ strongly in scutellar characteristics but are very different in pigmentation and pattern. A proposed history of the genus on Hispaniola suggests that there may be other populations of Diploglossus to be discovered there. As has been previously documented (Schwartz and Inchaustegui, 1976), the Antillean island of Hispaniola is a center of anguid lizard diversity. Four genera (Celestus Gray, Diploglossus Weigmann, Sauresia Gray, and Wet- morena Cochran) have representatives on Hispaniola (see Strahm and Schwartz, 1977, for a review of the genera of diploglossines); of these, Celestus has five species, Diploglossus one, Sauresia two, and Wetmorena one. According to Strahm and Schwartz’s analysis of the distributions of the genera of diploglossines on the mainland and in the West Indies, Dip- loglossus was the primary invader from the mainland, and spread northward from South America via the Lesser Antillean chain to Puerto Rico, Hispan- iola, and Cuba (but not Jamaica). On Puerto Rico and Cuba, there is only one species of Diploglossus; the species in each case are islandwide in distribution. On Hispaniola, there is also one species (D. warreni Schwartz); its distribution is peculiarly circumscribed in northern Haiti and on Ile de la Tortue. Celestus, on the other hand, reached Jamaica (where it is the only diploglossine genus and has had a broad species radiation) and His- paniola (where it has had a moderate radiation) from the Central American mainland and in addition gave rise to the elongate Sauresia with reduced limbs and the upland Wetmorena, which lacks external ear openings. Strahm and Schwartz (1977) attributed the limited range of D. warreni on Hispaniola to the facts that, despite its presumably early arrival there, the subsequent invasion and radiation of Celestus gradually forced Diploglossus to occupy only a portion of what might have been its former range. They alternatively suggested that perhaps Diploglossus has never had a broad distribution on Hispaniola, and that Celestus was successful because of the tenuous hold of Diploglossus on that island. It should be recalled that only on Hispaniola does more than one genus of diploglossines occur; even with- out reference to Sauresia or Wetmorena, no other island has both Celestus Dy PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON and Diploglossus. If this scenario has any validity, then it might be expected that other populations of Diploglossus exist in unexpected and isolated en- claves on Hispaniola, perhaps even far removed from northwestern Haiti. In the summer of 1977, Schwartz and Graham, in the company of Thomas M. Thurmond, spent 12 days in northern Haiti, basing themselves at Cap- Haitien. They were amazed at the fact that with little prompting local Hai- tians collected D. warreni in some numbers in the Vallée de Plaisance, within the rugged Massif du Nord. The result was the accumulation of 23 lizards from this region, three of which were kept alive. The elevation is about 336 m; the Plaisance area is extremely mesic. Although the species had been previously known from Limbé and its vicinity, some 12 km to the north, and from the Riviere des Barres, about 45 km to the northwest, the specimens from these two localities in collections number only five. Most startling was the taking of D. warreni at Terre Sonnain, 1.6 km N Les Poteaux, just east of Gonaives, in a relatively xeric situation. This single individual does not agree in coloration with the Limbé-Plaisance material, nor with that from Ile de la Tortue. Since we have but one specimen at this time, we are not willing to describe it as new, although we are convinced that it is a distinctive subspecies of D. warreni, Also available now are 15 specimens from Ie de la Tortue, many of which are juveniles. It appears that D. warreni, once considered a rather rare lizard with a limited distri- bution, is locally quite common and has a much broader northern Haitian distribution than previously conceived. In July 1977, the junior author, living in Santo Domingo as a Peace Corps Volunteer and working at the Parque Zoologico Nacional, sent the senior author a photograph of a large diploglossine which was one of four (two males, two females) taken 2.5 km west of Ingenio Catarey, at a place called ‘‘Come Hombre,”’ near Villa Altagracia, San Cristobal Province, Republica Dominicana. The lizards were collected by workmen clearing the area by bulldozer. The photograph (Fig. 1) of the living specimen showed that it was indeed very large, and Schwartz had little hesitation in calling it D. warreni, despite the gap of about 300 km between the Haitian and Domin- ican stations for the species. When the specimen died, it was brought to Schwartz. Although superficially the lizard is like D. warreni, it differs in a number of characters. The three other adults are still alive in the Parque Zoologico in Santo Domingo. Between 16 July and 3 August 1977, the two females gave birth to a total of 42 young. Of these, five died and were preserved, and have been examined by Schwartz and Graham. Some data have been taken by Duval from eight other living juveniles, as well as the three living adults in the Parque Zoologico. Some of the juveniles were distributed to the Bronx Zoo, the Philadelphia Zoo, and the Brookfield Zoo. Two of the Brookfield specimens have perished, and we have examined them in detail. Thus we VOLUME 92, NUMBER 1 3 sg Stns ape: ots pera c? a a oa .. ‘ : ‘ ‘ ee Fig. 1. Photograph of the holotype (USNM 197336) of D. anelpistus taken 22 June 1977 by Julian J. Duval. Note dark edging on head scales, lineate neck, and lack of sharp demarcation between dorsal and ventral colors. have complete data and counts on one adult and seven juveniles, as well as partial data on three adults and eight living juveniles. Diploglossus anelpistus, new species Holotype.—USNM 197336, adult male, from Ingenio Catarey, ‘‘Come Hombre,’’ 200 m, Villa Altagracia, San Cristobal Province, Republica Do- minicana, taken 21 July 1977 by Miguel A. Jardines. Paratypes.—ASFS V44945-46, ASFS V44949-51, ASFS V44954-55, ju- veniles born to females still living in the Parque Zoologico Nacional, be- tween 16 and 27 July 1977. Referred specimens.—Two females, alive at the Parque Zoologico at San- to Domingo, 22 and 23 June 1977; one male, alive at the Parque Zoologico Nacional, 13 July 1977; all paratypes and referred specimens are from the type-locality or born from females from the type-locality. Definition.—A species of Diploglossus closely related to D. warreni from northern Haiti (see Schwartz, 1970), but differing from that species in: (1) throat and venter very dark brown, the dark coloration on the chest and belly more or less involving individual scales, that on throat similar except the entire scales are dark with even darker centers; (2) all head scales out- lined with black; (3) nuchal scales posterior to enlarged head scales as far 4 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON as above auricular opening with dark pigment along their medial sutures to give a longitudinally striped neck; and (4) often with additional scales in- tercalated between the supralabials, especially between supralabials 3 and 4, or 4 and 5S. Description of holotype.—An adult male with a snout-vent length of 285 mm (all measurements in millimeters), tail length 112 (broken and only tip regenerated); weight prior to death 506.5 g; dorsum (as preserved) medium brown, the head not appreciably differently colored than remainder of body, the head scales all conspicuously outlined with black; the small nuchal scales likewise lined medially (but not posteriorly) with black to give nine longitudinal dark lines on the neck; 17 darker brown widely-opened chev- rons (=transverse bands), weakly angulate medially, each about 3 to 4 scales in width, on the back, these bands disappearing on the sides which are somewhat blotched, the scales becoming darker ventrally, so that between the chest and the vent the individual scales have dark brown centers and pale edges, the anterior scales with broader pale edges, the posterior ones with narrow or no pale edges; throat dark tan to medium brown, more or less uniform except that individual scales are still determinable by their pigmentation; limbs dark brown, unpatterned; 93 scales between mental and vent; 40 scales around body at midbody; angular subocular scale between supralabials 7 and 8 on one side, between 6 and 7 on the other; fourth toe lamellae 17; small intercalated scales (perhaps actually the uppermost por- tions of some of the supralabials) above supralabial series between supra- labials 3 and 4, and 4 and 5 bilaterally; head length 67.7, head width 49.0; arm length 58; longitudinal pale middorsal band 17 scales in width; head width/head length x 100 = 72.0. Variation.—Photographs of the three living adult specimens show that the venters in all three agree very well with that of the holotype. The ventral scales on the living specimens vary between 95 and 98. The male has 2 intercalated scales on the left side, between supralabials 2 and 3 and between 4 and 5, but none on the right side; one adult male has an intercalated scale above supralabial 4 on the right side but none on the left; the second female lacks intercalated scales. The juveniles (of which we have studied 7 preserved specimens and have data on 8 living ones) are patterned like the adults dorsally, but the ground color is much paler and the dorsal chevrons are much more bold. The dark edges of the large head scales are as apparent in the juveniles as in the adults. The venters of the juveniles are uniformly pale and lack the dark color of the adults. One preserved juvenile (ASFS V44949) shows nuchal streaking but this is less clear in the other preserved juveniles, where the neck has a more or less random dark pattern on a pale ground. One of the preserved juveniles has an intercalated scale between supralabials 3 and 4 on one side and none on the other; another has intercalated scales between VOLUME 92, NUMBER 1 5 supralabials 3 and 4, and 4 and 5, bilaterally; a third has an intercalated scale between supralabials 4 and 5 unilaterally; and a fourth has intercalated scales between supralabials 2 and 3, and 3 and 4 on one side, and between 3 and 4 on the other. Three juveniles lack intercalated scales. Of the eight living juveniles, six lack intercalated scales; one living specimen has an intercalated scale between supralabials 3 and 4 unilaterally, and the other (which has a congenitally deformed head), between supralabials 2 and 3 unilaterally. Thus, intercalated scales occur in 6 of 15 juveniles and in 3 of 4 adults. Since one of the females that contributed to the total of 42 young lacks intercalated scales, it is possible that all her offspring also lack them, accounting for the absence of such scales in 60% of the 15 juveniles studied. Both adult males have at least one intercalated scale unilaterally. Scale counts on 11 specimens (adult male holotype and 7 juveniles with com- plete counts; | adult male and 2 adult females with ventral counts only) are: scales between mental and vent 78—98 (x = 92.3); midbody scales 34-40 &@ = 37.0); position of angular subocular (by side) between supralabials 5/6 (2), 6/7 (13), 7/8 (1); fourth toe lamellae 16-19 @ = 16.9); dorsal band width 14- 18 scales @ = 16.0); number of dorsal chevrons 14-18 @ = 16.2); chin shields always 9. Comparisons.—The holotype of D. anelpistus is the largest Hispaniolan speciment of Diploglossus known; the snout-vent length of 285 exceeds that of the largest known D. warreni (ASFS V40338), which has a snout-vent length of 279. Likewise the head iength (67.2) and head width (49.0) are greater than the largest D. warreni (59.2 and 45.4, both from ASFS V40338). Comparative data between D. warreni (from both the Hispaniolan main- land and Ile de la Tortue) and D. anelpistus are shown in Table 1. None of the differences between the two species is striking or shows a strong di- chotomy between the two species. One scale relationship is important, but not absolute—no D. warreni has dorsally intercalated scales in the supra- labial series, whereas this is a common condition in D. anelpistus. It is in color and pattern that the two species differ most radically. Dorsal ground color in mainland D. warreni varies from deep brown to pale tan or gray; the ground color may have a metallic bronzy sheen. The chevrons are very dark brown to black, their intensity correlated with the depth of the ground color. The head shields are never outlined with black, and the head ground color may be deep russet (recorded in three adult males with snout- vent lengths between 210 and 279) or dull brownish orange (female, snout- vent length 260). The venters are never dark brown; rather, they are some shade of orange, from pale to deep. The chin and throat scales may be outlined with black or dark brown along their sutures, and there may be a few dark-edged scales randomly scattered on the belly, but the transition from the dark dorsal scales to the orange ventral scales follows a relatively sharp longitudinal division along the lower sides at the level of the limb 6 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Table 1. Mensural and meristic data for three populations of Hispaniolan Diploglossus. Position of angular subocular is tabulated by sides; although each specimen contributes double data for this figure, in some specimens one side may be injured and uncountable. Figures in parentheses are means. D. warreni D. warreni D. anelpistus (northern Haiti) (Tortue) Number 11 M5) 15 Largest (mm) Male 285 PUTS) 235 Female — 260 227 Scales, mental 78-98 (92.3) 82-97 (91.6) 81-98 (87.5) to vent Midbody scales 34-40 (37.0) 35-39 (37.4) 33-38 (35.6) Position of 5/6—2 /6—2 5/64 angular subocular 6/7—-13 6/7-43 6/7—20 7/8-1 7/8-4 7/81 4th toe lamellae 16-19 (16.9) 13-18 (15.7) 13=—15 (14.2) Dorsal band width 14-18 (16.0) 12-20 (16.6) 10-15 (13.9) Number of 14-18 (16.3) 15-19 (16.6) 14-17 (15.9) dorsal crossbands insertions. The most consistently dark scales are those along the margin of the anal flap; these scales may be solid dark brown for up to one-half of their length. The lower sides of D. warreni are more or less mottled with pale and dark tans and browns, rather than the more uniform dark condition in D. anelpistus. We have examined an osteoderm from D. anelpistus and find that it does not differ from osteoderms of D. warreni (Strahm and Schwartz, 1977). Remarks.—That the population of Diploglossus from Villa Altagracia is distinct from Haitian D. warreni is certain; pigmentation and pattern details are strikingly different. But there remains the question of the taxonomic status of the former; i.e., should it be regarded as a subspecies of D. warreni or aS a Species distinct from it? There seems little possibility of genetic continuity between the two populations, separated by 300 km from each other. Although it is likely that both populations are remnants of a formerly widespread species or species-complex, they seem now to be truly isolated. This has prompted us to name the Dominican lizards as a new species, rather than as a subspecies of D. warreni. Scutellar differences are slight, but pigmental and pattern differences are striking. It should be recalled that three of the Hispaniolan species of Celestus (stenurus Cope, costatus Cope, and curtissi Grant) differ little from one another in scutellation (Schwartz, 1964, 1970) but are easily separable by one familiar with the differences VOLUME 92, NUMBER 1 al between them. We emphasize this to demonstrate that scale differences within at least Hispaniolan diploglossines have not always evolved to a diagnostic level; more often, pattern and coloration play more important parts in species recognition than scale counts and scale relationships. When the junior author and his wife visited the type-locality on 12 July 1977, the bulldozer crew had nearly finished clearing. Less than 1.5 hectares remained of ‘‘Come Hombre.’’ According to a local resident, the forested area was about | km long and up to .25 km wide before clearing began. It was bordered on the west by treeless hills used as cattle pasture, and on the east by sugarcane fields. The section that the Duvals visited also had a small creek that ran parallel to the hills. The dominant tree seems to have been ‘“‘jabilla’’ (Hura crepitans), some with trunks nearly 1 m in diameter. Bromeliads (Jillandsia sp.) were common on the branches of these trees, along with some orchids (Epidendrum sp.). Philodendrum sp. and Dieffen- bachia sp. were also present in the small area that the crew had yet to clear. An important fact concerning the ‘‘Come Hombre’’ site is that it no longer exists as habitat. On 7 April 1978, the junior author visited the type-locality and found it planted in sugarcane. Some of the arroyos in the adjacent hills still have a few patches of trees, but otherwise the ‘‘Come Hombre’’ forest is gone. i The workers had unearthed the adult male that is still living in the Parque Zoologico in the root system of one of the few remaining “‘jabilla’’ trees, as the bulldozer knocked it down. The Duvals were told that this was the manner in which the other three adults had been collected, as well as several others that the workers had killed. Although the living specimens in Santo Domingo seem to be most active at night (which would make it easier for them to go unnoticed in the wild), it is noteworthy that the crew and a few local campesinos claimed never to have seen the lizard before the clearing began. In an effort to ascertain more information about these “‘/ucias grando- tas,’ as D. anelpistus is known locally, Duval asked residents of the area about them. Few claimed to be acquainted with the lizards, and some of those that did he found difficult to believe. In the settlement of Carvajal at 260 m, about 5 km to the southwest of ‘‘Come Hombre,”’ he spoke with two men who may be acquainted with D. anelpistus. One of them told Duval that someone had shot two or three of the lizards about two months earlier near the ‘‘Colmado del Ciego,’’ which is reached by following the first major creek south of ‘“‘Come Hombre’’ into the hills. The site was directly across the creek from the colmado. It is a north-facing overgrown (large trees) talus slope, planted with some young coffee trees; the area is less than 2 hectares in size. Duval spent about an hour on this slope and saw (but did not catch) one diploglossine about 160 mm long which may have been a young D. anelpistus. 8 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON If our previously noted scenario of the Diploglossus-Celestus history on Hispaniola is correct, it seems likely that other isolated populations of Dip- loglossus will be discovered. The fact that the type-locality of D. anelpistus is close (40 km) to Santo Domingo suggests that one need not look for other Diploglossus only in remote and little-known areas. The fact that four adults were collected (and others were killed) between 21 June and 13 July suggests that D. anelpistus was not especially rare in the vicinity of the type-locality. Similarly, D. warreni is not rare in northern Haiti; although we received 23 specimens (remarkably, all adults) in four days, we had the impression that the local population was virtually inexhaustible. The same may have been true of D. anelpistus. One should recall that after a single D. warreni was collected by W. L. Abbott at Riviere des Barres in 1917, no further speci- mens were taken until C. Rhea Warren secured them on Ile de la Tortue in 1968—a lapse of half a century. We call attention once again to Etheridge’s (1965) mention of two size-classes of fossil Diploglossus from a cave at Cerro de San Francisco near Banica, La Estrelleta Province, Republica Dominicana. The larger of these he estimated as being lizards with snout- vent lengths between 210 and 250 mm, precisely the size range of Hispan- iolan Diploglossus and much larger than any other known living Hispan- iolan diploglossine. The senior author has heard rumors of the presence of a large anguid in the Sierra de Neiba, but no specimens have been obtained. Finally, in May 1978, a large Diploglossus was collected at Comedero, between Los Conucos Adentro and Uniico, La Isabela, Puerto Plata Prov- ince, some 200 km north-northeast of the type-locality of D. anelpistus. All these facts tend to suggest that there remain isolated populations of Diplo- glossus on Hispaniola, and that we are just beginning to sample them. Etymology.—The name anelpistus is derived from the Greek for “‘unex- pected.”’ Specimens examined.—D. warreni: Haiti, Ile de la Tortue, near Palmiste (AMNH 103214, ASFS V15071); no other locality data (ASFS V20383, ASFS V23918, ASFS V20249-52, ASFS V20384-86, LDO 7-6422-23, LDO 7-6655-56); Dépt. du Nord, Riviere des Barres (USNM 59435); near Limbé (ASFS V26902, ASFS V26954, ASFS V26956, LDO 8-9101); 3.5 km S Plais- ance, 336 m (ASFS V40338-41, ASFS V40348-51, ASFS V40402-05, ASFS V40420-25, ASFS V44934, ASFS V45969); Dépt. de l’Artibonite, Terre Son- nain, 1.6 km N Les Poteaux, 122 m (ASFS V40243). Acknowledgments We are grateful to Thomas M. Thurmond for assistance in the field in Haiti and to Juan Berlino Bencosme and Miguel A. Jardines who brought three of the four adult specimens to the attention of the junior author. For previous loans of specimens we thank George R. Zug (National Museum of Natural History—USNM) and Richard G. Zweifel (American Museum of VOLUME 92, NUMBER 1 9 Natural History—AMNH). Lewis D. Ober kindly lent us specimens of D. warreni in his collection (LDO). Peter J. Tolson donated a specimen of D. warreni to the senior author, and animal dealer Louis W. Porras likewise has assured that, upon their deaths, living specimens that have passed through his hands would be returned and given to the senior author (Albert Schwartz Field Series—ASFS). Larry D. Wilson was instrumental in bring- ing the holotype of D. anelpistus to the senior author. Ray Pawley of the Brookfield Zoo kindly sent us two juveniles that had died in that collection. The staff of Ingenio Catarey helped the junior author locate ‘‘Come Hombre’’ and’allowed him to accompany the clearing crew on 13 July 1977. We are also grateful to ZOODOM and the Peace Corps, Republica Do- minicana, for their support. To all of the above we express our gratitude, since without their help and assistance, we would have far less material of these large and showy lizards. Literature Cited Etheridge, R. E. 1965. Fossil lizards from the Dominican Republic.—Q.J. Florida Acad. Sci. 28(1):83-105. Schwartz, A. 1964. Diploglossus costatus Cope (Sauria, Anguidae) and its relatives in His- paniola.—Reading Public Mus. and Art Gallery, Sci. Publ. 13:1-—57. . 1970. A new species of large Diploglossus (Sauria: Anguidae) from Hispaniola.— Proc. Biol. Soc, Wash. 82(60):777-788. , and S. J. Inchaustegui. 1976. A new species of Diploglossus (Reptilia, Lacertilia, Anguidae) from Hispaniola.—J. Herpetology 19(3):241-246. Strahm, M. H., and A. Schwartz. 1977. Osteoderms in the anguid lizard subfamily Diplo- glossinae and their taxonomic importance.—Biotropica 9(1):58-72. Miami-Dade Community College North, Miami, FL 33167; 717 SW Ist St., Hallandale, FL 33009; and Parque Zooldgico Nacional, ZOODOM, Apartado Postal 2449, Santo Domingo, Republica Dominicana. PROC. BIOL. SOC. WASH. 92(1), 1979, pp. 10-22 SOME PHYLOGENETIC IMPLICATIONS OF A DISCOVERY OF ASPIDORAS PAUCIRADIATUS (PISCES: SILURIFORMES: CALLICHTHYIDAE) FROM THE RIO NEGRO IN BRAZIL Stanley H. Weitzman and Martha H. Balph Abstract.—Aspidoras pauciradiatus is redescribed on the basis of 10 specimens collected from the Rio Negro near Tapurucuara, Amazonas, Bra- zil. The occurrence of Aspidoras nearly 3,000 river kilometers beyond the previously known range of the genus brings into question statements by H. Nissen and I. J. H. Isbrucker concerning the distribution and relationships of Aspidoras and Corydoras. Substantial variability in size of the frontal cranial frontanel may invalidate the use of this character to separate Aspi- doras from Corydoras, although the two genera still may be distinct on the basis of the supraoccipital fossa. Additional collecting is needed to test further the hypothesis that Aspidoras and Corydoras should be recognized as separate genera. Aspidoras pauciradiatus (Weitzman and Nijssen) has been known from two type specimens collected in 1961 from the Rio Araguaia near Aruana, State of Goias, Brazil (Weitzman and Nijssen, 1970) and from three aquar- ium specimens without known locality (Nijssen and Isbrticker, 1976). The objectives of this paper are: (1) to report on what may be 10 additional specimens of this species (Figs. 1—3) collected in 1972 from the Rio Negro near Tapurucuara, State of Amazonas, Brazil, over 3,000 river kilometers from the type locality and nearly that far out of the previously known range of the genus Aspidoras von thering; and (2) to discuss relationships between populations of A. pauciradiatus and between Aspidoras and Corydoras Lacépede in the light of this discovery. Aspidoras pauciradiatus (Weitzman and Nijssen) Figs. 1-3, Table 1 Synonymy.—See Nijssen and Isbrticker (1976: 114). Specimens examined.—1, holotype, National Museum of Natural Histo- ry, United States National Museum (USNM) 191625, SL 23.2 mm, Brazil, State of Goias, Rio Araguaia, near Aruana (14°58’'S, 51°04’W), H. R. Ax- elrod, 1960. 1, paratype, USNM 204363, SL 22.6 mm, same data as holo- type. Ten distributed in museums as follows: 2, Academy of Natural Sci- ences, Philadelphia (ANSP) 136687; 2, British Museum (Natural History) VOLUME 92, NUMBER 1 11 Fig. 1. Aspidoras pauciradiatus, 6MZUSP, SL 16.5 mm, Brazil, State of Amazonas, Rio Negro, Sao Joao, near Tapurucuara. (BMNH) 1978. 18:1—2; 2, Museu de Zoologia da Universidade de Sao Paulo (MZUSP); 2, USNM :218375; 2, Zoological Museum of Amsterdam (ZMA) 115.205; SL 15.7-17.7 mm, Brazil, State of Amazonas, Rio Negro, Sao Joao, near Tapurucuara (00°24’S, 65°02'W), P. Vanzolini, 23 September 1972. Diagnosis.—A. pauciradiatus may be distinguished from all other species of Aspidoras and from ail species of Corydoras and Brochis Cope by the following combination of character states: dorsal fin I,6; anal fin I,5; pectoral fin 1,7; medial border of pectoral-fin spine prominently serrated with 10-15 large spinules (Weitzman and Nijssen, 1970:129, Fig. 6); lateral body scutes 23/20; supraoccipital shield with a small, circular fossa on its central dorsal surface. Description.—Morphometric and meristic data are in Table 1. We present measurements of more parameters than Nijssen (1970) suggests, because we ate Fig. 2. Aspidoras pauciradiatus, 6MZUSP, SL 16.5 mm, Brazil, State of Amazonas, Rio Negro, Sao Joao, near Tapurucuara. WW PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON ee ae Fig. 3. Aspidoras pauciradiatus, 2MZUSP, SL 17.4 mm, Brazil, State of Amazonas, Rio Negro, Sao Joao, near Tapurucuara. believe these measurements may show useful interspecific and interpopu- lational differences and similarities in Aspidoras, Corydoras, and Brochis. Body rather heavyset. Snout blunt, rounded at tip. Two rictal and one mandibular barbel on each side of head. Small, bony prickles present on surfaces of all scutes, bones, and fin spines but not on belly or soft fin rays. Color in alcohol.—Color of the holotype and paratype was described by Weitzman and Nijssen (1970). The Rio Negro specimens are described be- low. Basic (‘‘ground’’) body and head color pale brown; markings brown to grayish brown, probably nearly black in life. Top of head very dark, nearly black in some specimens; no dark mask in area of eye. Dorsum of snout anterior to interorbital area paler than dorsum of head and with a pale brown band extending between posterior nares. Anterior to pale band, dorsum of snout mottled with more or less triangular or vermiform dark marks. Just ventral to nares, an irregular, nearly horizontal pale brown stripe extends from near snout tip posteriorly to anterior border of eye. Area ventral to this pale region with a broad dark brown stripe extending from snout tip and origin of dorsal rictal barbel posteriorly and dorsally to anterior border of eye. Often entire and always basal portion of dorsal rictal barbel dark brown. Ventral rictal barbel pale brown posterior to base, which is shared with dorsal rictal barbel; distal portion of ventral rictal barbel with scattered dark brown chromatophores. Mandibular barbel with a patch of dark chro- matophores at base; free portion of mandibular barbel pale brown with scattered dark chromatophores. Area ventral and posterior to eye nearly white with a rim of dark pigment near border of eye. Opercle mostly with interconnecting dark brown blotches of moderate size; ventral border of opercle nearly white. Ventral portion of head, especially ventral area of branchiostegal rays and isthmus, without dark pigment patches or dark chro- matophores. Mandibular region nearly white but bearing scattered dark chromatophores. Dorsolateral area of branchiostegal rays with scattered VOLUME 92, NUMBER 1 13 dark chromatophores just ventral to opercle. Dorsum of body and supra- occipital region of head dark brown, often mottled with paler brown; dark brown pigment often confined to posterior borders of head plates and dor- solateral body scutes. External surface of cleithrum and coracoid bones heavily mottled with rather evenly spaced patches of closely set dark brown chromatophores. Sides of body with dark and very pale patches arranged as a Series of clear to rather vague posteriorly pointing chevrons with apices at shared junctions of dorso- and ventrolateral body scutes. Belly without dark pigment from head posteriorly to area of pelvic fins and anus. Dorsal fin with a broad, somewhat horizontal, nearly black stripe at or near its base, this dark pigment often missing or much paler on proximal anterior portion of dorsal-fin spine and at or near base of posteriormost fin rays. Distal to this dark stripe, dorsal-fin rays and membrane nearly white except for a more or less horizontal, very dark stripe extending from near distal tip of spine across distal portions of anterior three or four fin rays. Distal tips of spine and anterior fin rays white; distal tip of first fin ray sometimes black. Tips of middle fin rays may be black and distal third or half of posterior three rays white. Spine of adipose fin darkly pigmented, white at tip; remainder of adipose fin mostly black except at base. Caudal fin with four, five, or six irregular black vertical bars; pigment entirely or almost entirely confined to fin rays. Distal portions of circular body scutes at base of each caudal-fin lobe black. Anal fin pale except for three or four dark markings extending vertically across fin rays. Pelvic fins hyaline except for two or three broad, irregular rows of black bars across fin rays. Pectoral fins with four or five broad rows of dark bars on fin rays; anterior spine dark, especially where rows of fin-ray pigment extend onto spine. Color in life. —A photograph in Axelrod et al. (1967:F 223.00) shows color in life of the paratype. Pigmentation of living Rio Negro specimens has not been observed. Discussion Relationship between Rio Araguaia and Rio Negro populations of Aspi- doras pauciradiatus.—Specimens from the Rio Araguaia and from the Rio Negro resemble each other in many ways. These admittedly small samples do not differ in a majority of morphometric and meristic characters (Table 1). They also have nearly identical color patterns, as may be seen by com- pairing Figs. 1-3 with that in Nijssen and Isbrticker (1976:115, Fig. 5). Both are small forms relative to most species of Aspidoras and Corydoras; all specimens examined seemed to be sexually mature, and females contained eggs, at 16-23 mm in standard length. Fishes of both populations exhibit the following diagnostic combination of character states, defined for A. pauci- radiatus by Weitzman and Nijssen (1970) or by Nijssen and Isbriicker PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 14 SN 6c I tac a) o5C = 00 0C ¢S9JNds uy-osodipesid snosAze jo 19qUINN — a = 00 OT =? 0°0 Ol $9}Nds uy-[esiopsid snosAze jo 1oquinN SN cl ¢°L6-S°L8 ce L'16 9°06-L°L8 OC C68 e4isus] sulds uy-[e10}59g SN 8r 1 ¢°LE—-0'8¢ 9¢ Tee i WEG OC cl c0t 4isua] ouids uy-ssodipy x 8L°C 0°98-0'FL a 4 v 6L 6 IL-7 69 61 DYE e4isua] sulds uy-jesiog x voll L’y9-0°9S 9C Gals 6 96-S°L8 we) (O69) pelOQieg [BJO [eQUIA JO YISUIT SN 80°1 € 98-0 PL 6€ v 08 $°L8-0°08 es 88 e4IPIM peoy jsoyeosH SN el 0 Sec-P SI VC c 6! 0:0c 8 81 80 v6! coved jesiopoid Jo yysus] ek vy v CPc-S 81 [El il IZ, 6 9I-¥'6 es¢ Gell pe[SURIUOJ [BIUBIO [eIUOIJ JO YSUIT SN Id] 86-0P OC (ee to) LAM) te te bs eS[PUIQIOeIFUL Je1jUsA Jo yJdop jsoyeoIH ek Done 0 VE—-9'6C bil 9°CE C 6C— 1 8¢ 80 9°8¢ eHIQIO JO 1OJOWRIP JSIJBoIT) SN vS'l 0'0S—-S er IG 9°9F eSp-l ey Sy Ii (nA e[B11G10.19}UT AUOG JO YIPIM IsvIT SN O10 L’Ly-0'8t [L, © 6° CY Sty-S' Iv yl 4 4idus] NOUS x tye €tIl-v 6 OT Oa ) tell 17 toll 0 Y) fe zSPplOov105 JO soseqg JOLI9}Ue UZdMJOq JDURISIG ek 9t'P OLI-I v1 80 9'SI yeI-9'Cl 9°0 Oe! z2[ounped jepned jo yidap isvo'] SN v0°0 9 1€-8°8C 80 €0¢ 8° 0£—6 62 9°0 vOt zyidep Apog jsaye2a1D *k 9V'¢ 6 8S—-C SS (aa TLS BSS-LES | 8 Ps zo ue JO 1aps1og 10L19}Ue 0} dy ynous SN e1'0 y 16S €8 vc L98 S°L8-9°98 9°0 0-48 zUISII0 UY-ssodipe 0} diy ynoug SN €0'I 6° €8-$'8L cl p18 0'i8-S"6L I'l Z'08 zUISIIO UY-[eue 0} dij jnoUs kk Sle O1S-S°Lh GI 0 6P ¥ 90-6 SP v0 c OF zUISLIO UY-[esIOp 0} dy jnoug x LOE SC 1€—-8°8¢ 80 c Ot (O) (OE 1G 60 v8? z4)su2] peoH x 60 CI [ELM =IE, SII L0 L9l A? 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Although the Rio Araguaia and Rio Negro specimens exhibit a number of similarities, they differ in certain respects. In Rio Araguaia specimens the rictal barbels extend beyond the eye to reach the posterior border of the opercle, whereas in Rio Negro specimens these barbels do not reach the posterior border of the eye. The frontal cranial fontanel is circular to oval in outline in the Rio Araguaia fishes but is elongate and oval in those from the Rio Negro. Dark pigment is absent from the pectoral and pelvic fins of Rio Araguaia specimens but is present on those of Rio Negro specimens. Finally, the two samples show statistically significant differences in 11 of 31 morphometric and meristic characters (Table 1). The significant differences between the Rio Araguaia and Rio Negro spec- imens most likely reflect genetic differences between the two populations, although certain alternative explanations are possible and the samples are small. If the differences are hereditary, we cannot at present determine whether the Rio Araguaia and Rio Negro specimens represent different species or geographical variants of a single species, as collections of small fishes from intervening localities are fragmentary. Future sampling in such areas may indicate whether or not morphologically intermediate forms exist and thus help to resolve this issue. In the meantime, we think it best to act conservatively by classifying the Rio Negro specimens as A. pauciradiatus. Relationship between Aspidoras and Corydoras.—Nissen and Isbrucker (1976) characterize Aspidoras by its possession of two cranial fontanels— a frontal fontanel that is shared with other callichthyid genera and a supra- occipital fontanel (fossa) unique to Aspidoras. They further note that the frontal fontanel is much smaller and more circular in outline in Aspidoras than in the closely related genus Corydoras. They found the structure of the skull roof to be essentially similar in 400 specimens representing 13 species of Aspidoras and observed ‘‘no intermediate structure’’ (1976: 109) in more than 100 species of Corydoras. Our recent discovery of specimens presumed to be A. pauciradiatus from the Rio Negro, as well as a reevaluation of some earlier material, obscures in part the taxonomic distinction between Aspidoras and Corydoras. The Rio Negro fishes did possess a frontal cranial fontanel and a supraoccipital fossa, but the frontal fontanel was significantly longer than in Rio Araguaia specimens (Table 1). Furthermore, individual variation, particularly within the Rio Araguaia sample, was considerable. The holotype from the Rio Araguaia exhibits a small, nearly circular frontal fontanel, similar to other species of Aspidoras as illustrated by Nijssen and Isbrticker (1976: 126, Fig. 16), whereas the paratype from the same locality has a relatively elongate frontal fontanel that more closely resembles those of the Rio Negro speci- VOLUME 92, NUMBER 1 17 mens and certain species of Corydoras. One could speculate that an en- larged frontal fontanel is paedomorphic for A. pauciradiatus, associated with its pygmyism and therefore progenic, although the interpopulational and individual variation that we observed in adults of this species might not support this interpretation. Adults of large species of Aspidoras and espe- cially Corydoras show a considerable amount of interspecific variation in frontal fontanel size. Significant intraspecific differences in the size of the frontal fontanel in adults of A. pauciradiatus, as well as substantial inter- specific variation in this character in Corydoras, causes us to question the use of the size or shape of the frontal fontanel to distinguish Aspidoras from Corydoras, as the structure of this fontanel may be more labile than was previously supposed. The supraoccipital fontanel or fossa appears to be found only in Aspidoras among adults of the callichthyid genera, as pointed out by Nijssen and Isbrucker (1976:109). We could find no evidence of an isolated supraocci- pital fossa in adult or juvenile specimens of C. pygmaeus Knaack (USNM 218355, 5 specimens, SL 10.2—-18.1 mm); C. aeneus Gill (ZMA, 2 specimens, SL 10.6-10.7 mm); C., c.f. caqguetae Fowler (USNM 218358, 130 specimens, SL 7.3-36.9 mm); C. elegans Steindachner (USNM 218359, 105 specimens, SL 21.2-33.0 mm); C. barbatus Jenyns (USNM 100916, 6 specimens, SL 36.8-70.7 mm); and C. eques Steindachner (SU-CAS 17746, 10 specimens, SL 7.8-33.5 mm). Among most of the Corydoras adults that we examined, the frontal fon- tanel bifurcated only the anterior tip of the supraoccipital; however, in ju- veniles it extended posteriorly through the supraoccipital almost to the bone’s midpoint, the region where the supraoccipital fossa occurs in species of Aspidoras. In adults of C. elegans (USNM 218359) the frontal fontanel bifurcated the anterior portion of the supraoccipital bone into right and left halves. These observations suggest that the supraoccipital fossa of Aspi- doras may have evolved as an isolated section of a frontal fontanel that originally extended deeply into the supraoccipital. It further seems possible that the supraoccipital fossa is neotenic for large species of Aspidoras but progenic for the pygmy species A. pauciradiatus. If such is the case, it is conceivable that the supraoccipital fossa is independently derived (and hence non-homologous) in pygmy and non-pygmy species of Aspidoras, a circumstance which could lessen the utility of this character for separating Aspidoras from Corydoras. Further problems are posed by the discovery of a population of a species of Aspidoras almost 3,000 river kilometers out of the known range of the genus (Fig. 4). Previously, the genus Aspidoras was known only from a fairly restricted area in southern and eastern Brazil (including the Parana basin, a few eastern coastal drainages to the north and south of the Rio Sao Francisco, and southern portions of the Rio Tocantins and the Rio Xingu); 18 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON { ie \ i b c= A. PN : Fee \ = ZS NS TA i ds = | Nw 4 ; ry SD Pe 1S —S a =e Fig. 4. Distribution of Aspidoras based upon Nijssen and Isbriicker (1976:130, Figs. 20- 21), with the addition of the Rio Negro locality reported here for A. pauciradiatus. Circle enclosing star represents type locality of A. pauciradiatus, black circle represents Rio Negro locality of same, and black dots represent localities of other species of Aspidoras. it was believed never to have occupied the main Amazon basin (Nijssen and Isbrucker, 1976). Conversely, Corydoras was known to be widely distrib- uted in the Orinoco and Amazon basins, the Guianas, the eastern highlands of Brazil, many of the eastern coastal streams of Brazil, and the Parana basin in Paraguay (Fig. 5). On the basis of these ostensible differences in distribution, Nissen and Isbrticker (1976) concluded that Corydoras might be more advanced than Aspidoras if the presumptive ancestor of both gen- VOLUME 92, NUMBER 1 19 7 al D> = » = ae = e \\ et w 1G SP oe ew. Ss m—~ if AS = ee Fig. 5. Distribution of Corydoras based upon reports in the literature. Doubtful or non- specific localities have been excluded. Although Nijssen (1970:59, Fig. 35) indicates Corydoras from the Rio Magdalena in Colombia, this locality may be in error (e.g. see Eigenmann, 1923:65, 220, 228; Miles, 1947:100). era was restricted to southern and eastern Brazil. That specimens of A. pauciradiatus have now been found in the upper central portion of the Amazon basin brings these authors’ statements concerning the distribution and relationships of Aspidoras and Corydoras into question. Our discussion to this point is based upon an assumption that the Rio Negro specimens do in fact represent A. pauciradiatus. If they do not, their bearing upon the relationship of Aspidoras and Corydoras may or may not 20 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON differ from what we have described. For instance, the Rio Negro specimens might represent a new species that is closely related to A. pauciradiatus. If this is true, our interpopulational comparisons become interspecific com- parisons, but our remarks concerning the relationship of Aspidoras to Co- rydoras still apply. Alternatively, the similarities between the Rio Araguaia and Rio Negro specimens might be due in part to convergence in two species of Aspidoras that are not closely related. Once again, the size and shape of the frontal fontanel in the Rio Negro specimens raise questions concerning the appropriateness of using this character to separate Aspidoras from Co- rydoras. The possibility also remains that the enlarged frontal fontanel and/ or the supraoccipital fossa are progenic for the new species of Aspidoras. Finally, it is conceivable that the Rio Negro specimens represent a new, aberrant, pygmy species of Corydoras which exhibits a supraoccipital fossa independently derived by progenesis and in a cladistic sense not homologous with that found in Aspidoras. If such is the case, the discovery of these specimens far out of the known range of the genus Aspidoras no longer poses problems. We think it unlikely that similarities between the Rio Araguaia and Rio Negro specimens are due to convergence of unrelated forms. Certain shared characters, such as pygmyism, reduced number of dorsal- and pectoral-fin rays, and reduced number of dorso- and ventrolateral body scutes, appear important in this regard. Pygmyism is not known elsewhere in Aspidoras but apparently has occurred at least twice in Corydoras, once in the pre- sumably related species C. pygmaeus and C. hastatus Eigenmann and in- dependently in C. habrosus Weitzman. Except for small adult size, how- ever, we can find no shared derived character to relate the Rio Araguaia and Rio Negro specimens to pygmy species of Corydoras; hence we think it likely that their small adult size is independently derived. The Rio Ara- guaia and Rio Negro specimens also exhibit a shared color pattern not found in other species of Aspidoras or Corydoras, although it is difficult to assess the usefulness of this character as a test of relationships. Since, on balance, a few apparently derived characters are shared only by the Rio Araguaia and Rio Negro specimens, we believe that the two forms are closely related. Although the derivation of the supraoccipital fossa in these pygmy forms could be independent of that in large species of Aspidoras, we suspect that the supraoccipital fossa probably is homologous in all forms that exhibit it. At this time we do not have sufficient information to make an objective judgment concerning the relationship of Aspidoras to Corydoras. Despite questions raised by the discovery of the Rio Negro specimens, we tend to favor the idea that Aspidoras is phylogenetically distinct from Corydoras on the basis of the supraoccipital fossa. We also suspect that Aspidoras and Corydoras may differ in head shape and body shape, although we do not have access to sufficient material to examine this possibility at present. VOLUME 92, NUMBER 1 21 The number and distributional patterns of recorded collections of Aspi- doras and Corydoras indicate that fishes of these genera have not yet been adequately sampled. Additional collecting efforts are needed to test further the hypothesis that Aspidoras and Corydoras should be recognized as sep- arate genera. Acknowledgments Paulo Vanzolini (MZUSP) graciously allowed the senior author to survey and borrow specimens from Amazonian fish collections accumulated under his direction. These collections are supported by the Fundacao de Amparo a Pesquisa do Estado de Sao Paulo, which since 1967 has maintained the Expedicao Permanente da Amazonia. Heraldo Britski and Naercio Menezes (MZUSP) provided invaluable aid in locating specimens and locality infor- mation. Marilyn Weitzman, William Fink, and Sara Fink assisted in the survey of collections at the MZUSP. Travel funds were provided by the Smithsonian Institution Amazonian Ecosystems Program, directed by Clif- ford Evans. Resumo.—Este trabalho refere-se a descoberta de 10 exemplares tenta- tivamente identificados como Aspidoras pauciradiatus, coletados no Rio Negro, proximo de Tapurucuara, Amazonas, Brasil, que se situa aproxi- madamente 3.000 quilometros fora da area de distribuigao conhecida do genero Aspidoras. A. pauciradiatus € redescrita com base nos exemplares do Rio Negro. Embora esses exemplares apresentem diferengas signifi- cantes (P 0.05) em relacao aos exemplares-tipo do Rio Araguaia em 11 dos 31 caracteres meristicos e morfometricos estudados, nos acreditamos que as duas amostras devem ser consideradas co-especificas até que se disponha de material coletado nas localidades intermediarias para comparacoes. Se os exemplares do Rio Negro nao sao A. pauciradiatus, € possivel que eles reprentem uma nova espécie intimamente relacionada a esta espécie. A ocorréncia de Aspidoras na parte central superior da Bacia Amazonica demonstra que as consideracgoes feitas por Nijssen e Isbrucker (1976) sobre a distribuigao de Aspidoras e Corydoras e sobre as relagoes entre esses géneros precisam ser realiadas. A consideravel variacgao interindividual e interpopulacional do tamanho da fontanela craniana frontal nos adultos de A. pauciradiatus, bem como a variabilidade interespecifica deste carater em Aspidoras e particularmente em Corydoras, leva-nos a questionar 0 uso do tamanho da fontanela craniana frontal para separar Aspidoras de Corydo- ras. Os dois géneros pode ainda ser distinguidos com base na fossa supra- occipital, embora seja admissivel que esta estrutura tenha surgido indepen- dentemente em diferentes espécies, sendo por isso nao homologa no sentido 22 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON cladistico. Assim, o reconhecimento de Aspidoras como um “‘taxon’’ di- ferente de Corydoras pode ou nao ser uma medida apropriada. Coletas posteriores poderao ajudar a resolver estes problemas. Literature Cited Axelrod, H. R., C. W. Emmens, D. Sculthorpe, W. Vorderwinkler, and N. Pronek. 1967. Exotic tropical fishes: M 3.00-121.00, P. 2.00—93.00, C 3.00—48.00, F 1.00-608.00.—T. F. H. Publications, Jersey City. Eigenmann, C. H. 1923. The fishes of western South America, I—Mem. Carnegie Mus. 9(1): 1-346. Miles, C. 1947. Los peces del Rio Magdalena.—Ministerio de la Economia Nacional, Colom- bia, 1-xxviil plus 214 pp. Nijssen, H. 1970. Revision of the Surinam catfishes of the genus Corydoras Lacépede, 1803 (Pisces, Siluriformes, Callichthyidae).—Beaufortia 18(230): 1-75. , and I. J. H. Isbriicker. 1976. The South American plated catfish genus Aspidoras R. von Ihering, 1907, with descriptions of nine new species from Brazil (Pisces, Siluri- formes, Callichthyidae)—Bijdragen Tot de Dierkunde 46(1):107-131. Weitzman, S. H., and H. Nijssen. 1970. Four new species and one new subspecies of the catfish genus Corydoras from Ecuador, Colombia and Brazil (Pisces, Siluriformes, Cal- lichthyidae).—Beaufortia 18(233):119-132. (SHW) Department of Vertebrate Zoology, Smithsonian Institution, Washington, D.C. 20560; (MHB) Department of Wildlife Science, Utah State University, Logan, Utah 84322. PROC. BIOL. SOC. WASH. 92(1), 1979, pp. 23-27 THE HUMERUS OF XENICIBIS, THE EXTINCT FLIGHTLESS IBIS OF JAMAICA Storrs L. Olson and David W. Steadman Abstract.—The first known complete humerus of the extinct fossil ibis Xenicibis xympithecus is reported from a new locality in Jamaica. The spec- imen exhibits many of the morphological characteristics seen in flightless birds and possesses a number of characters which distinguish it readily from all other ibises. At the time of our description of the extinct ibis Xenicibis xympithecus (Olson and Steadman, 1977) from a Quaternary deposit in Long Mile Cave, Trelawny Parish, Jamaica, a fragmentary shaft of a humerus was the only element of the wing available. In this, part of the distal end of the deltoid crest was preserved, and its morphology, along with that of the paratypical coracoids, led us to the conclusion that X. xympithecus was flightless. Shortly after reading proofs of our paper, we were apprised of an unfamiliar humerus among a collection of Jamaican fossils at the University of Florida. Upon comparing this specimen with the paratypical humeral fragment (AMNH 11031), we found the same diagnostic twisting and flattening of the shaft in both and, except for the paratype being slightly more robust, the two are similar in all other details as well. As there is hardly any possibility of there being two different species of flightless birds in Jamaica of the same size and with the same details of the humerus, we conclude that the new specimen must pertain to Xenicibis xympithecus and it is therefore of im- portance in further elucidating the morphology of the wing of this bizarre form. The specimen (Figs. | and 2) is a complete and beautifully preserved right humerus (Florida State Museum UF 23768) collected from Swansea Cave I (Hole 3, L3) near Worthy Park, St. Catherine Parish, Jamaica, by T. H. Patton in June 1966. This locality is near the center of the island and lies about 57 airline km ESE of the type locality. McGrath (1960) provides a map and general description of Swansea Cave, but makes no mention of any fossiliferous deposits. The age of the deposit in question is not known and we can only assume it to be Quaternary. As with the type material of X. xympithecus, we compared this specimen with a synoptic series of humeri of Recent non-passerine families of birds, finding it to be most similar to ibises (Threskiornithidae). Although in a number of ways unique, this humerus shows several distinctive characters 24 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 1. Right humeri in palmar view. a, Apteribis glenos, B. P. Bishop Museum BBM-X 147240; b, Xenicibis xympithecus, Florida State Museum UF 23768; c, Eudocimus albus, USNM 500883. All figures natural size. which in combination indicate certainly that the bird formerly possessing it was flightless: slender twisted shaft, very deep brachial depression, bicipital crest reduced, deltoid crest reduced in area, thickened, and twisted; external tuberosity reduced and displaced distally. The new specimen is longer than the humerus of Eudocimus, Plegadis, or Mesembrinibis, but is shorter than that of Threskiornis, Harpiprion, Theristicus, Hagedashia, or Platalea. Because elements of the hindlimb of Xenicibis are larger than in any of these genera, the size of the humerus must be small relative to the size of the bird, as would be expected in a flightless species. The measurements of the bone are as follows: overall length 112.6 mm, width of proximal end 23.0, depth of head 7.8, length of deltoid crest from VOLUME 92, NUMBER I 25 Fig. 2. a—d, right humerus of Xenicibis xympithecus, UF 23768; a, palmar-internal view; b, internal view; c, external view; d, anconal view. e, paratypical fragment of shaft of right humerus of Xenicibis xympithecus, AMNH 11031. All figures natural size. proximal surface of external tuberosity 27.0, least width of shaft 6.2, width of shaft at midpoint 7.1, depth of shaft at midpoint 5.8, width of distal end 16.2, depth through external condyle 10.0, greatest diameter of brachial depression 9.8. Compared to extant genera of ibises, the humerus in Xenicibis has the shaft much more slender and greatly curved both latero-medially and dorso- ventrally. The shaft is twisted and becomes peculiarly flattened in the por- tion beginning between the deltoid crest and the midpoint of the shaft and continuing distally for about 30 mm. In anconal view the head is large and deep, extending much farther distally than in other genera. The capital groove is much deeper, longer and more distinct. Beneath the internal tub- 26 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON erosity is a single small, oval, well-defined pneumatic foramen, unlike the ill-defined and more cancellous pneumatic foramina of extant ibises. The external tuberosity is reduced, forms a much steeper diagonal slope with the head, and is marked by a distinct pit on the anconal surface. The center of the anconal surface of the shaft just distal to the head and the internal tuberosity is markedly concave. The point of attachment of M. latissimus dorsi posterioris is marked by a distinct oval pit, 3.2 mm long, whereas in all other ibises this is a raised prominence. The bicipital crest and attach- ment for M. scapulohumeralis caudalis are greatly reduced; in extant ibises the bicipital crest extends distally well beyond the level of the internal tub- erosity, whereas in Xenicibis the area of the internal tuberosity in anconal view nearly obscures the bicipital area. The deltoid crest in Xenicibis is small but thickened, with the area of the pectoral attachment bearing a protuberance. At the distal end, the brachial depression in Xenicibis is much more extensive and much deeper than in any other known ibis, occupying most of the width of the shaft. Likewise, there is a much deeper depression between the attachment of the anterior articular ligament and the internal condyle. In external view the attachment of the anterior articular ligament is more reduced and more distally situated, and the external condyle pro- trudes more palmarly than in other ibises. The olecranal fossa is deeper in its distal portion, while the entepicondyle is less elevated anconally above the tricipital grooves than in extant ibises. The only other flightless ibis discovered is Apteribis glenos Olson and Wetmore 1976, from the islands of Molokai and Maui in Hawaii. The hu- merus of this form (Fig. la) shows many of the same modifications as that of Xenicibis but differs greatly in other respects as follows: shaft more robust, rounded, and much less curved; head more elongate, without distal projection; pneumatic foramen absent; external tuberosity without pit on anconal surface; internal tuberosity with long axis nearly perpendicular to shaft, not parallel; deltoid crest more swollen; attachment of M. latissimus dorsi posterioris not concave; entepicondylar area more pointed, projecting farther internally; brachial depression much smaller and shallower. Unfortunately, the humerus of Xenicibis is so greatly modified that it sheds no light on the relationships of this enigmatic genus. The new locality record reported here suggests that X. xympithecus was once widespread in Jamaica. Its remains can be expected in cave deposits elsewhere on the island. Acknowledgments For calling the new specimen of Xenicibis to our attention we are indebted to Gary S. Morgan. For his comments on the manuscript we are grateful to John Farrand, Jr. The photographs are by Victor E. Krantz. VOLUME 92, NUMBER 1 27 Literature Cited McGrath, B. R. G. 1960. Swansea Cave, St. Catherine.—Geonotes 3(1):23-26. Olson, S. L., and D. W. Steadman. 1977. A new genus of flightless ibis (Threskiornithidae) and other fossil birds from cave deposits in Jamaica.—Proc. Biol. Soc. Wash. 90(2):447— 457. Olson, S. L., and A. Wetmore. 1976. Preliminary diagnoses of two extraordinary new genera of birds from Pleistocene deposits in the Hawaiian Islands.—Proc. Biol. Soc. Wash. 89(18):247-258. (SLO) Department of Vertebrate Zoology, National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560; (DWS) Depart- ment of Geosciences, University of Arizona, Tucson, Arizona 85721. PROC. BIOL. SOC. WASH. 92(1), 1979, pp. 28-41 REVIEW OF TOXOTHERIUM (PERISSODACTYLA: RHINOCEROTOIDEA) WITH NEW MATERIAL FROM THE EARLY OLIGOCENE OF WYOMING Robert J. Emry Abstract.—Previously unreported specimens of the diminutive rhinocer- otoid, Toxotherium, from the White River Formation (early Chadronian Age) of central Wyoming add new anatomical information and confirm some previous interpretations based on less nearly complete material. Occurring in a single thin stratigraphic interval are permanent lower dentitions refer- able to Toxotherium hunteri, deciduous lower dentitions referable to T. woodi, and upper dentitions referable to Schizotheroides jackwilsoni. The latter two species are placed in the synonymy of T. hunteri. Toxotherium is most likely an amynodontid. Introduction During the past ten years, several specimens of the rare, diminutive rhinocerotoid, Toxotherium, have accumulated as a result of collecting in the White River Formation of the Flagstaff Rim area in central Wyoming (Emry, 1973). Only four specimens of Toxotherium have been reported previously, each from a different locality, ranging from southern Saskatchewan to Trans-Pecos Texas. The new material includes maxillary dentition believed to represent Toxotherium, and also provides information suggesting that T. woodi is based on a juvenile 7. hunteri and is therefore a synonym of T. hunteri. The upper dentition believed to be 7. hunteri represents the same taxon as Schizotheroides jackwilsoni Schiebout, 1977, which is also placed in the synonymy of 7. hunteri. Abbreviations for the following institutional collections are used: AMNH, The American Museum of Natural History, New York; F:AM, Frick Col- lection, American Museum of Natural History; NMC, National Museum of Canada, Ottawa; SDSM, South Dakota School of Mines and Technology, Rapid City; TMM, Texas Memorial Museum, University of Texas, Austin; USNM, National Museum of Natural History, Smithsonian Institution, Washington, D.C. VOLUME 92, NUMBER 1 29 Order Perissodactyla Suborder Ceratomorpha Superfamily Rhinocerotoidea Family Incertae sedis Genus Toxotherium Wood, 1961 Toxotherium hunteri Wood, 1961. Toxotherium woodi Skinner and Gooris, 1966, new synonymy. Schizotheroides jackwilsoni Schiebout, 1977, new synonymy. Type.—NMC 8918, anterior half of right mandibular ramus with P.-M,. Hypodigm.—Type and: F:AM 42901 (Type of T. woodi), partial left man- dibular ramus with dP,(broken)-dP,,M,; TMM 40206-22, left M?; TMM 40840-42, maxillary fragment with right P*-M? (type of Schizotheroides jack- wilsoni); TMM 40283-100, anterior part of right mandicular ramus with dP, ; SDSM 8442, a partial left mandibular ramus with P,-M,; USNM 244352, left mandibular ramus with dP.4, M,; USNM 244353, part of right M,; USNM 244354, right dP;; USNM 244355, left dP.; USNM 244356, left dP,; USNM 244357, left dP;; USNM 244358, right M'; USNM 244359, right maxillary fragment with M'?; USNM 244360, posterior two-thirds of left M,; USNM 244361, left mandibular fragment with P.-M,; F:AM 79403, right dP.,. Known distribution.—Southern Saskatchewan, central Wyoming, eastern Wyoming, and Trans-Pecos Texas. The type of Toxotherium hunteri is from the Chadronian Cypress Hills Formation of Saskatchewan. Toxotherium was next reported from central Wyoming by Skinner and Gooris (1966), who described the new species T. woodi, which I believe on the basis of present evidence is juvenile 7. hunteri, and which is from the lower part of the White River Formation, early Chadronian in age. Toxotherium was next recorded by Harris (1967) in the Ash Springs Local Fauna of the undiffer- entiated Vieja Group of the Sierra Vieja area of Trans-Pecos Texas, an occurrence also regarded as early Chadronian. Emry (1973) recorded Tox- otherium in the fauna of the White River Formation in the Flagstaff Rim area of central Wyoming on the basis of a single tooth (F: AM 79403). SDSM 8442 is from the Yoder Local Fauna, early Chadronian, of Goshen Hole Wyoming (see Kihm, 1975). The two specimens described by Schiebout (1977) as Schizotheroides jackwilsoni, are, I believe, the upper dentitions of Toxotherium hunteri, for reasons explained below. All the other speci- mens listed in the hypodigm above (i.e., all the specimens with USNM numbers and F:AM 79403) are from the lower (early Chadronian) part of the White River Formation in the Flagstaff Rim area of central Wyoming. Of these, USNM 244352 through 244357 and F: AM 79403 came from a single rich concentration of bone that has many other mammalian taxa. This quarry is near the head of Little Lone Tree Gulch, at 44 feet below ash B on the 30 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON generalized zonation section of the Flagstaff Rim sequence (Emry, 1973, p. 29). The other specimens, USNM 244358 through 244361, came from a small area along the south side of Little Lone Tree Gulch, in the vicinity of the quarry, and within a relatively thin stratigraphic interval, the lowest occur- rence at approximately 50 feet below ash B, and the highest at about 5-10 feet below ash B. The local range-zone for Toxotherium in the Flagstaff Rim section is from about 120 to 170 feet on the generalized section (Emry, 1973), although all but one specimen occurred in the much thinner interval between about 120 and 130 feet. The White River Formation here is more than 700 feet thick. The occurrence of Toxotherium only in this relatively restricted interval in a much thicker section suggests either that it had a very short temporal range, or that it preferred some particular environmen- tal conditions that existed in the area only during the time in which this thin interval was being deposited. All of the known occurrences of Toxotherium are Chadronian, and all for which more precise information is available (i.e., all except for the type of T. hunteri) are known to be early Chadronian. Although Toxotherium is now known from enough localities to define a rather large geographic range, it was apparently never an abundant element of any Chadronian fauna. Wood (1961) suggested that the late discovery of Toxotherium, and its ap- parent rarity, might be explained by its being a northern form, with the southern limit of its range near the Cypress Hills of southern Saskatchewan. The specimen from southern Saskatchewan now defines the northernmost limit of the known range, which extends southward as far as Trans-Pecos Texas. Associated fauna.—The new material of Toxotherium is associated with a large number of vertebrate taxa, most of which are recognized as typical of Chadronian Age (though not necessarily restricted to the Chadronian), and some of which are indicative of early Chadronian. Among the taxa occurring in the quarry concentration with Toxotherium are the following: Peratherium, Oligoryctes cf. O. altitalonidus, Leptictid cf. ‘‘Ictops’’ acu- tidens, Apternodus, Ischyromys (or Titanotheriomys), Protosciurus, Cylin- drodon, cf. Yoderimys, Adjidaumo, Paradjidaumo, ?Namatomys, Auloli- thomys, Heliscomys, Palaeolagus, Daphoenocyon, Hesperocyon, ?Parictis, Hyracodon, Merycoidodon, Agriochoerus, Poebrotherium, Hypisodus, and the samples of Leptomeryx discussed by Emry (1973) as species B and D. — Fig. 1. Toxotherium hunteri, USNM 244352, left mandibular ramus of young individual with dP,.,, M,. A, occlusal view, stereogram, B, lateral view. Approximately x2, scale in millimeters. Compare with F:AM 42901 (Skinner and Gooris, 1966, fig. 4, type of T. woodi), which is at same stage of ontogenetic development. 31 VOLUME 92, NUMBER 1 A a LA jy PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Dental formula of Toxotherium.—Wood (1961) identified the teeth pres- ent in the type of 7. hunteri as P».4, with P, preceded by alveoli for P,. This interpretation was questioned by Skinner and Gooris (1966), who argued that the teeth present in the specimen are P,-M, with alveoli for P;. Skinner and Gooris (1966) identified the most posterior tooth preserved in F:AM 42901 (type of 7. woodi) as M, because of its ‘‘abrupt increase in size and less wear on the crown relative to the ‘preceding teeth.’’ This interpretation is valid if the preceding teeth are correctly interpreted as deciduous pre- molars, an interpretation with which I agree, and for which additional evi- dence is found in the new material, particularly in USNM 244352 (Fig. 1). This jaw is virtually a duplicate of F: AM 42901, except that the most anterior tooth is complete rather than broken. The preservation is typical of the White River Formation, with pale tan to cream colored bone and brown teeth. In fossil mammals with this kind of preservation, and in which the deciduous dentition can be positively distinguished from the permanent, the color of the enamel of the deciduous dentition is usually distinctly lighter than that of the permanent teeth. The reason for this color difference has not been demonstrated, to my knowledge, although it is presumably related either to the relative thickness of the enamel (thinner in deciduous than in permanent teeth) or to some difference in its internal structure. In the Tox- otherium jaw from Little Lone Tree Gulch, the anterior three teeth are distinctly lighter in color than the posterior one, indicating that the teeth are dP..4, M,, as Skinner and Gooris had determined for the type of T. woodi. The color difference is not apparent in Figure | because the specimen was given a light coat of ammonium chloride before it was photographed. I interpret the teeth of the Vieja specimen (TMM 40283-100) as dP,s, rather than P;., as Harris (1967) believed them to be. Harris stated that the dimensions of the teeth of his specimen agree well with those of dP.., of T. woodi, and stated also that he was following the interpretation of Skinner and Gooris. The morphology of the more posterior of the two teeth is cer- tainly more like that of dP; of T. woodi than is it like that of the P, of T. hunteri. Too little is known about the maxilla of Toxotherium to allow even an educated guess as to the formula of the maxillary dentition. On the basis of present knowledge, the formula for the lower deciduous cheek teeth is dP,.4, and for the permanent dentition is P34, M,-3. This requires the as- sumption that dP, is not replaced by a permanent Pz, but it is not unusual among mammals for the most anterior deciduous premolar not to be re- placed, and this phenomenon seems to be more prevalent among groups undergoing reduction of the premolar series. The ‘‘tusk’’ of Toxotherium.—Wood (1961), Skinner and Gooris (1966), Harris (1967), and Radinsky (1969), all commented on the large lower front tooth of Toxotherium, although in only one specimen (F:AM 42901, type of T. woodi) is even part of the tooth known. In the other specimens the VOLUME 92, NUMBER 1 33 ‘‘disproportionately large’’ size of the tooth is an interpretation based on part of the outer wall and the basal part of its alveolus. Wood (1961), for example, as part of the diagnosis of Toxotherium, stated that it has a ‘‘sin- gle, disproportionately large, bulbous-rooted lower front tooth (incisor or canine).’’ The type of T. woodi has part of the tooth (called a tusk, to avoid the problem of homology) in place in the alveolus. Skinner and Gooris (1966) noted that the enamel of the tusk in their specimen ‘‘extends well back but not to the extreme base of the root,’ and that the root is ‘“‘open, a condition usually reflecting prolonged growth.’’ Any interpretation of the relative size of this tusk, however, must consider the age of the individual, which was very young (with the M, erupted but essentially unworn, and M, not yet erupted). I interpret the tusk of this specimen as an unerupted permanent caniniform tooth, with the root not yet formed. The “‘disproportionately large’’ size of the tooth can be accounted for, at least in part, by its being a permanent tooth only partly formed in an immature jaw. Skinner and Gooris (1966) mentioned that the cross section of the ‘‘tusk”’ is about 41% of the ramal depth, but this comparison has little relevance. The depth of the jaw at this point is through the symphyseal area where the jaw is shallowest, and in this immature specimen the depth of the jaw at any point will be less than might be expected in a mature individual, whereas the diameter of the crown of the tusk would not increase once it had formed. A more significant comparison might be between the size of the tusk and the size of the cheek teeth. According to Skinner and Gooris, the greatest diameter of the ‘‘tusk at break’’ is 5.8 mm, or less than the anteroposterior dimension of dP,, which is 6 mm. If the tusk is correctly interpreted as a permanent tooth, it would be even more meaningful to compare its size with that of a permanent cheek tooth. In this specimen (F:AM 42901) the only permanent cheek tooth present is M,, which is 10 mm anteroposteriorly, according to Skinner and Gooris. The greatest cross-sectional diameter of the tusk is, then, 58% of the greatest dimension of M,. The tusk does not seem disproportionately large in this context. To determine how the tusk size, relative to M,, of Toxotherium compares with that of other rhinocerotoids, a number of similar measurements were made on specimens in USNM collections. In the discussion following, tusk measurements are of the greatest cross-sectional dimension, and the M, measurements are of the greatest anteroposterior dimension; all are in mm. In two specimens of Trigonias the tusks are 16.4 and 20.6, while the re- spective first molars are 34.5 and 32.2. In five specimens of Subhyracodon, the tusks are 19.7, 20.4, 20.0, 23.1, and 18.2, while the respective first molars are 29.9, 29.8, 29.6, 31.5, and 30.1. The tusk (second incisor in rhinocero- tids) measurement averages 55% of the M, measurement in Trigonias and 67% in Subhyracodon. In two specimens of Metamynodon in the USNM collections, the tusks (which in amynodonts are canines) are 37.5 and 47.4 in their largest cross- 34 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON sectional dimension, while the respective first molars are 42.1 and 47.3. In these two specimens the tusk measurement averages 95% of the first molar measurement, though this statistic masks one instance in which the tusk measurement slightly exceeds the first molar measurement. These statistics show that the tusk of Toxotherium, relative to the first molar, is not disproportionately large when compared to those of other rhinocerotoids. It is relatively slightly larger than that of Trigonias, some- what smaller than that of Subhyracodon, and much smaller than that of Metamynodon. In terms of tusk size, Toxotherium is not an unusual rhin- ocerotoid. Identity of Toxotherium woodi.—In the introduction of their paper de- scribing Toxotherium woodi, Skinner and Gooris (1966) stated that ‘‘certain characters separate the Bates Hole specimen, specifically at least, from T. hunteri.’’ They did not provide a diagnosis for the species, however, and no distinguishing characters were specifically mentioned elsewhere in the text except for the statement (1966:9) that in 7. woodi the dental series is ‘shortened by the loss of P,, not by both P, and P, as in 7. hunteri.’’ This conclusion assumes that the deciduous P, of T. woodi is replaced by a permanent P,, an assumption for which there is no evidence. The material discovered since that time does not clarify this question. The permanent dentitions known either have no P.,, or are too incomplete to determine whether or not P, was present. All the juvenile specimens with anterior premolars preserved have dP,. The type of 7. hunteri is a jaw of an individual that was still young but approaching maturity, with M, slightly worn and M; not yet erupted. The type of 7. woodi represents a much younger individual, with the deciduous premolars still in place, M, practically unworn, and M, not yet erupted. Dental comparisons between the two specimens are therefore limited to M,, | which, as noted by Skinner and Gooris (1966), are morphologically similar | and very nearly the same size in the two specimens. While 7. hunteri ap- pears to be considerably larger, the size difference is surely related to dif- — ferent ontogenetic development; dentally the size difference is insignificant | (Table 1). The new material from central Wyoming provides additional information suggesting that T. woodi represents a young individual of T. hunteri. USNM 244352 (Fig. 1) is virtually a duplicate of the type of T. woodi, except that — dP, is more nearly complete. The other deciduous teeth from the same quarry (see hypodigm above) are morphologically like those of the type of T. woodi. There would be no hesitation in assigning all the juvenile speci- mens to T. woodi, were it not for the fact that in the very same thin strati- | graphic interval in the same very restricted area, specimens with permanent dentition occur that could be assigned without hesitation to T. Aunteri. USNM 244361 (Fig. 2), a jaw fragment bearing P,-M,, compares very well | VOLUME 92, NUMBER 1 35 Table 1. Measurements, in millimeters, of teeth of Toxotherium hunteri. Specimen Maximum length, Maximum width, number Tooth anteroposterior transverse TMM 40206-22 M? 16.3 18.1 TMM 40840-42 ee 9.4 (minimum) 11.8 M! 13e7 13.9 M? 18.2 17.5 (estimate) M? igez 20.2 USNM 244359 M! 1233 i22 M? 14.8 15.0 (estimate) USNM 244358 M! 11.5 ial NMC 8918 (type) P.-M, 32.6 PE 8.3 5.6 M, 10.3 6.1 M, 14.1 8.0 F:AM 42901 dP,-M, 26.6 dP, 728 4.6 dP, 8.8 Sys M, 10.0 6.3 USNM 244352 dP.-M, 28.2 dP3-M, 252 dP, 3.6 25 dP; dae 4.2 dP, 8.3 5.1 M, 9.9 6.2 USNM 244361 len JES 5.6 M, 9.4 G2, TMM 40283-100 dP, 5.0 3.0 dP; 7.8 4.5 SDSM 8442 Ry 8.3 S65) M, 10.2 6.4 USNM 244355 dP, 4.3 p26 USNM 244354 dP; 622 3.8 USNM 244356 dP, 7.6 4.0 USNM 244357 dP, Fike 4.2 F:AM 79403 dP, 8.6 4.8 USNM 244353 M, 6.1 USNM 244360 M, 7.9 with the type of T. hunteri; it is slightly smaller but certainly within the range of variation that might be expected in a single species. The depth of the jaw of this specimen cannot be determined because the lower border is not preserved, but even the fragment remaining is much deeper than the entire jaw below M, of USNM 244352 (Fig. 1) which has a slightly larger M, but is a much younger individual. The maxillary fragment with M?? (USNM 244359, Fig. 3) at first appears to be too large to be Toxotherium, _ but when it is occluded with the type of T. hunteri (the only lower jaw of 36 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 2. Toxotherium hunteri, USNM 244361, left mandibular fragment with P.-M,. A, oc- clusal view, stereogram; B, lateral view. Approximately x2, scale in millimeters. Compare with NMC 8918 (Wood, 1961, figs. 1-3, type of T. hunteri). Toxotherium with an M, and M, to occlude) it is seen to be precisely the right size. The wear facets of the upper teeth all occlude correctly with those of the lowers. These upper teeth, and an isolated M' (USNM 244358), were found in the same thin stratigraphic interval in which the lower teeth and jaws of Toxotherium occur. Neither upper dentitions like those men- tioned nor lowers of Toxotherium occur elsewhere in the more than 700 feet of White River section. In the interval in which they occur, there are no other upper dentitions that could conceivably be associated with Toxoth- erium and no other lower dentitions that could conceivably be associated with these upper dentitions. Though the association is circumstantial, I think | | | | VOLUME 92, NUMBER 1 37 wii Fig. 3. Toxotherium hunteri, USNM 244359, right maxillary fragment with M' and most of M?, stereogram. Approximately x2, scale in millimeters. Compare with TMM 40840-42 (Schie- bout, 1977, fig. 1f, type of Schizotheroides jackwilsoni). the evidence is quite persuasive that these upper dentitions are of Toxo- therium. WIS, The lack of any morphological features to separate T. hunteri and T. woodi, and the association of specimens assignable to both species in the same deposits, leads to the conclusion that 7. woodi merely represents very young individuals of T. hunteri. I therefore place T. woodi in the synonymy of T. hunteri. Identity of Schizotheroides jackwilsoni.—Skinner and Gooris (1966) dis- counted the idea that Toxotherium might represent the lower dentition of Schizotheroides Hough (1955), an enigmatic perissodactyl known only from the late Eocene of Sage Creek, Montana. Their reasons were that the cusp pattern of Schizotheroides is not as distinctly rhinocerotoid as that of Tox- otherium, and they believed that upper dentition with heavy cingula, as in Schizotheroides, was not likely to be combined with lower dentition with very weak, or no, cingula, as in Toxotherium. Schiebout (1977), in her description of what she regarded as a new species of Schizotheroides, cited the reasons given by Skinner and Gooris for not associating Schizotheroides and Toxotherium. Here the problem begins, because the material on which Schiebout based Schizotheroides jackwilsoni certainly represents the same taxon as the upper dentitions from Little Lone Tree Gulch (USNM 244358 and 244359), which I believe, for reasons discussed above, represent the upper dentition of Toxotherium hunteri. I believe that the Vieja specimens are also Toxotherium, but not that Toxotherium is a synonym of Schizo- theroides. The two specimens from the Vieja Group occur in local faunas in which 38 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON lower dentitions of Toxotherium have not been found, but Toxotherium is recorded (Harris, 1967) in the Vieja Group in the nearby Ash Springs Local Fauna. As in the White River Formation, there are no other upper dentitions that could be associated with Toxotherium, and no other lower dentitions that could be associated with Schizotheroides jackwilsoni. Though the association can be proven only by finding upper dentitions articulated, or directly associated, with lowers, I believe the present evi- dence is persuasive. As pointed out above, the association of upper and lower dentitions in the Flagstaff Rim sequence is even more compelling. I therefore place Schizotheroides jackwilsoni in the synonymy of Toxother- ium hunteri. As Schiebout (1977, p. 456) mentioned, S. jackwilsoni is larger and more hypsodont than S. parvus, the genotypic species, and in S. jackwilsoni the third molar is both longer and wider than the second, whereas the opposite is true of S. parvus. There are other differences not mentioned by Schie- bout. In S. parvus, the protolophs turn posteriorly at the lingual ends, most noticeably on M?, so that these lophs are concave posteriorly when seen in occlusal view, in contrast to those of the Vieja and Little Lone Tree Gulch specimens in which the protolophs and metalophs are concave anteriorly, particularly on M! and M?. S. parvus has a continuous cingulum from the parastyle across the anterior, lingual, and posterior border, to the posterior end of the metacone, on both M? and M®? (Schiebout’s illustration of S. parvus does not show a cingulum around the lingual end of the protoloph of M?®, but it is present on the specimen, and shows clearly in Hough’s original illustration, 1955, pl. 8, fig. 1). S. parvus has a distinct labial cin- gulum, more prominent on M? than on M?, and on M? a small but distinct mesostyle. This is in contrast to the Vieja and Little Lone Tree Gulch specimens which have a distinct anterior cingulum which ends at the lingual end of the protoloph, no labial or lingual cingula on M! and M?, and a very short and weak posterior cingulum, barely indicated on M’ and M?. The third molars illustrated by Schiebout (1977, fig. 1d-f) have weak lingual cingula, and one has a faint suggestion of a labial cingulum. None has a mesostyle. Schiebout’s illustration of §. parvus (1977, fig. la—b) is inaccurate in several details which are clearly shown in Hough’s original photographic illustration of the specimen (1955, pl. 8, fig. 1). The cingulum of M? was mentioned above. Schiebout’s illustration shows no hint of a protoconule, which is quite distinct in the specimen, particularly on M?, and in Hough’s photograph. The paracone is much more distinct than shown by Schiebout; the protocone, particularly of M?, turns posteriorly much more than indi- cated by Schiebout; and the valley separating the paracone and parastyle is incorrectly placed in Schiebout’s drawing. This valley, which is shown by Schiebout as intersecting the ectoloph posterior to the protoloph-ecto- VOLUME 92, NUMBER 1 39 loph junction, is actually at the junction, opposite the valley separating the protoloph and parastyle, so that the ectoloph, protoloph, and parastyle meet at a common point. The large parastyle of M?, with its crest oriented anterolabially-postero- lingually, is the one unusual feature common to Schizotheroides parvus and the upper dentition here referred to Toxotherium hunteri. While this may suggest some relationship between the two taxa, the many differences point- ed out above are, in my opinion, sufficient to separate them at least at the generic level. Not enough is known, particularly of S. parvus, to make a good case either for or against a relationship. Toxotherium hunteri is cer- tainly more derived than S. parvus ina number of features, such as complete merging of the protoconule into the protoloph, increased hypsodonty, and loss of lingual, labial, and most of the posterior cingula of the molars. Discussion.—Toxotherium has resisted placement in a taxonomic cate- gory lower than superfamily. Wood (1961) was apparently confident that it belongs to the Suborder Ceratomorpha, and believed that it is probably assignable to the Superfamily Rhinocerotoidea. Skinner and Gooris (1966) assigned Toxotherium to the Rhinocerotoidea, as have the other authors who have since mentioned it. Radinsky (1969) noted that Toxotherium has a rhinocerotoid molar pattern, and an enlarged, procumbent tusk, and sug- gested that it may be an aberrant, diminutive amynodontid. The new ma- terial of Toxotherium does little to resolve the matter, but the additional information is more suggestive of amynodont affinities than any other al- ternative. Amynodonts typically have massive skulls and jaws, with the facial part of the skull relatively short for a perissodactyl. Toxotherium was very small, but relative to the size of its teeth the dentary was massive, and the rela- tively wide angle between the lower jaws (mentioned by Skinner and Gooris, 1966, and Harris, 1967) suggests that Toxotherium was brachycephalic. Oth- er trends in amynodonts (Radinsky, 1969) are reduction of the premolar Series, increasing hypsodonty and relative narrowing of the molars, with Some increase in obliquity of the cross crests. These trends are also manifest in Toxotherium, which has lost both P, and P, of its permanent dentition. The teeth are relatively high crowned for a rhinocerotoid with so little mo- larization of the premolars, a condition noted by Wood (1961), and the cross crests of the molars are somewhat more oblique than in rhinocerotids and much more oblique than in tapiroids. In the upper dentition (Fig. 3, and Schiebout 1977, figure 1f), the cross lophs are also oriented at an oblique angle to the ectoloph. The upper dentition assigned here to Toxotherium would be somewhat contradictory as an amynodont because of the large parastyle of M?. Typical amynodonts have parastyles that are considerably smaller, more columnar, and not deflected anterobuccally. But this parastyle is unusual for any pe- 40 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON rissodactyl and would be a contradictory feature in assigning Toxotherium to any family. Schiebout (1977) did point out that, despite the unusual form of the parastyle in the Vieja specimens, it nevertheless functions just as it does in the hyracodontid rhinocerotoid Triplopus, by forming, along with the posterior end of the ectoloph of the next anterior tooth, a surface which shears against the labial side of the protoconid. Because the evidence is so limited and in part contradictory, it seems best for the present to defer assigning Toxotherium to a family, and simply to leave it in Rhinocerotoidea incertae sedis. Schizotheroides, like Toxotherium, has resisted assignment to suprage- neric categories. Hough (1955) placed it questionably in the Superfamily Chalicotheroidea. Radinsky (1964) noted that the parastyle of Schizo- theroides parvus is more compressed anterolingually-posterolabially than in chalicotheres, and that ‘“‘the only other features at all suggestive of chali- cothere affinities are a slight swelling midway down the posterior side of the worn M? protoloph, and, also on M?’, a sharp cingular cuspule between paracone and metacone (which is probably merely an abnormal variation, but may indicate incipient mesostyle development).’’ Radinsky regarded the absence of a distinct protoconule as a strong argument against placing Schizotheroides in the Chalicotheroidea (1964). Although I agree with Ra- dinsky that Schizotheroides is not a chalicotheroid, it should be pointed out that Schizotheroides does have more distinct protoconules than Radinsky’s comments would suggest. It seems incongruous that Radinsky mentioned the trace of a protoconule on the worn M7?, but did not mention that M? has a protoconule even more distinct. Neither did Hough mention the presence of a protoconule in Schizotheroides parvus, though it is distinctly evident in the illustration accompanying her type description (Hough 1955, pl. 8, fig. 1). The protoconules of Schizotheroides parvus are more distinct than in any of the many Homogalax specimens in the USNM collections. Ra- dinsky (1969) noted that a characteristic of tapiroids is the complete lack of protoconules in the molars, except in Homogalax, the earliest known ta- piroid, which retains traces of protoconules. If the absence of protoconules is characteristic of any tapiroid later than Homogalax, then Schizotheroides is excluded from the Tapiroidea, since it has more distinct protoconules and occurs later than Homogalax. It is not an objective of this report to classify Schizotheroides; more material of S. parvus is needed before the question of its affinities can be answered. I will follow Radinsky (1964) in leaving it in Perissodactyla in- certae sedis. Acknowledgments Some of the most nearly complete specimens mentioned in this report were found by volunteer field assistants, who deserve special mention; the VOLUME 92, NUMBER | 4] jaw shown in Fig. | was found by Elizabeth Hunter and that shown in Fig. 2 was found by Jennifer Emry. The manuscript was read by John A. Wilson, Morris F. Skinner, Shirley M. Skinner, Clayton Ray, and Donald Prothero; I thank them for improvements resulting from their comments and correc- tions. Literature Cited Emry, R. J. 1973. Stratigraphy and preliminary biostratigraphy of the Flagstaff Rim area, Natrona County, Wyoming.—Smithsonian Contr. to Paleobiol. 18:1—43. Harris, J. M. 1967. Toxotherium (Mammalia: Rhinocerotoidea) from western Jeff Davis Coun- ty, Texas.—Pearce-Sellards Series 9: 1-7. Hough, J. 1955. An upper Eocene fauna from the Sage Creek area, Beaverhead County, Montana.—Jour. Paleont. 29:22—36, pls. 7-8. Kihm, A. J. 1975. Mammalian paleontology of the Yoder Local Fauna.—Unpublished masters thesis, South Dakota School of Mines and Technology, Rapid City, S. D. Radinsky, L. B. 1964. Paleomoropus, a new early Eocene chalicothere (Mammalia, Peris- sodactyla), and a revision of Eocene chalicotheres.—Amer. Mus. Novitates 2179: 1-28. . 1969. The early evolution of the Perissodactyla.—Evolution 23:308-—328. Schiebout, J. A. 1977. Schizotheroides (Mammalia: Perissodactyla) from the Oligocene of Trans-Pecos Texas.—Jour. Paleont. 51:455-458. Skinner, S. M., and R. J. Gooris. 1966. A note on Toxotherium (Mammalia, Rhinocerotoidea) from Natrona County, Wyoming.—Amer. Mus. Novitates 2261:1-—12. Wood, H. E., Il. 1961. Toxotherium hunteri, a peculiar new Oligocene mammal from Sas- katchewan.—Nat. Hist. Papers, Natl. Mus. Canada 13:1—-4. Department of Paleobiology, National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560. PROC. BIOL. SOC. WASH. 92(1), 1979, pp. 42-44 A NEW SPECIES OF LIMNODRILUS (OLIGOCHAETA: TUBIFICIDAE) FROM JAMAICA Ralph O. Brinkhurst Abstract.—The new tubificid oligochaete Limnodrilus variesetosus from Jamaica differs from the other twelve or thirteen species in the genus in the form of the setae and of the penis sheaths. Loden (1977) recently described two new Limnodrilus species from the southeastern United States, bringing the total list of species to 12 (or 13 if one includes L. spiralis as a recognizable species). While Loden suggested that L. grandisetosus be removed from synonymy with L. silvani in that paper, subsequent investigation has caused him to reverse that stand by virtue of evidence that he will present elsewhere. This evidence relates to the presence of giant ventral setae in specimens with penis sheaths of the silvani type, which is highly characteristic. In L. rubripenis the ventral setae of IV or V to IX differ from the rest, making only two species in which the dorsal and ventral setae are not always identical. Specimens sent to the author from Jamaica by M. C. Taylor (Fisheries and Environment Canada) proved to have ventral setae from H-IV or V different from the rest, but otherwise all of the characteristics of the genus Limnodrilus, and hence these must be regarded as belonging to a new species. Limnodrilus variesetosus new species Fig. | Holotype. —NMNH 5617. Jamaica: Black River, Holland, 23.3.78, below a sugar factory. | slide. Coll.E.Ross. Paratypes.—NMNH 56172. Same locality and date, 10 specimens in al- cohol. Etymology .—‘‘variesetosus’’—setae not all alike. Description.—Immature worms 12-14 mm x 0.3-—0.5 mm, mature worms up to 50 mm X 1.0 mm, segments up to 170. Ventral setae of II-IV or some of V 4-5 (immature) or 6-8 in number, generally most numerous in II, large with very long upper teeth. Dorsal anterior setae slightly fewer in number, upper teeth less elongate. All other setae diminishing to 2 or 3 per bundle progressively, teeth of about equal length, the lower thicker than the upper. Reproductive system with all structures paired and of characteristic form for the genus. Atria somewhat narrow and tubular; prostates attached near proximal end, glands very large and lobed. Penis sheaths about 4—5 times VOLUME 92, NUMBER 1 43 G 0-1 Fig. 1. Limnodrilus variesetosus. A, Ventral setae of IV; B, Dorsal setae of IV; C, Setae of median segment; D, Spermathecal vestibule and duct; E, Penis sheath; F, Atrium and pros- tate gland; G, Spermatozeugmata. All lengths in mm. longer than average breadth, basal third tapering quite sharply, the rest more or less cylindrical. Spermathecal ducts relatively long, terminating in vestibulae. Spermatozeugmata elongate, slightly expanded, club-like at one end. Discussion.—This is a typical Limnodrilus species, with bifid setae and long, heavily cuticularized, penis sheaths surrounded by spiral muscles. It is distinguishable from L. udekemianus in that the upper teeth of the anterior ventral setae are not significantly thicker than the lower, and those setae are limited to a few anterior ventral bundles. The penis sheaths of L. udek- emianus and the new species are basically similar, but differ in proportions and form from those of other species in the genus (Brinkhurst and Jamieson, OTA). 44 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Acknowledgments I wish to thank Dr. M. C. Taylor for the provision of specimens, and Mr. M. S. Loden for specimens of other species in the genus. Literature Cited Brinkhurst, R. O., and B. G. M. Jamieson. 1971. Aquatic Oligochaeta of the World.—Uni- versity of Toronto Press. Toronto. Loden, M. S. 1977. Two new species of Limnodrilus (Oligochaeta, Tubificidae) from the Southeastern United States ——Trans. Amer. Micros. Soc. 96(3):321-326. Institute of Ocean Sciences, P.O. Box 1700, Sidney, B.C. Canada V8L 382. Addendum: J. Madill (National Museum of Canada, Museum of Natural Sciences) has examined additional material from the same Jamaican source, and finds the dorsal setae closely resemble the ventrals, the elongate upper teeth present from IIJ-V or V. The distal end of the ventral setae may be more strongly curved than the dorsals, but this is uncertain. The apparent length of the setal teeth is strongly affected by orientation, of course, and most of Ms. Madill’s specimens are too poorly oriented to confirm the last point. PROC. BIOL. SOC. WASH. 92(1), 1979, pp. 45-50 REDESCRIPTION OF THE THECAL TABULATION OF SCRIPPSIELLA GREGARIA (LOMBARD AND CAPON) COMB. NOV. (PYRRHOPHYTA) WITH LIGHT AND SCANNING ELECTRON MICROSCOPY Alfred R. Loeblich, III, James L. Sherley, and Robert J. Schmidt Abstract.—Peridinium gregarium is transferred to Scrippsiella and the thecal tabulation is revised. Variations in this plate tabulation occur. The thecal formula most common is p.p., p.r., 4’, 3a, 6”, 6c, Ss, 5”, 2”. On 1 November 1967, A. Loeblich, III isolated a single cell from a water sample collected by Eugene Lombard from a tidepool in Palos Verdes Pen- insula, Los Angeles County, California. This isolation resulted in a clonal culture, UTEX 1948. A clone of the organism from the 1967 Lombard col- lection which was initially identified as Peridinium sociale was used in a study of lipids (Lee & Loeblich, 1971). Lombard & Capon (1971) described this same dinoflagellate as a new species, Peridinium gregarium Lombard & Capon, 1971. They believed P. gregarium to be closely related to Peri- dinium sociale (Henneguy ex Labbé) Biecheler, 1935, and figured (Fig. 2A, C) a tabulation that was identical to that given by Biecheler (1935) for P. sociale. The theca of Peridinium gregarium was stated to differ from that of P. sociale in 1) lacking an apical projection, 2) possessing an ‘‘antapical pore’’ (actually figured on a sulcal plate and not on an antapical plate), 3) having striated rather than reticulate ornamentations and 4) having dif- ferent thecal plate shapes. We examined the theca of isolate UTEX 1948 of P. gregarium using light and scanning electron microscopy. Our findings presented below differ sig- nificantly from the light microscopic observations of Lombard & Capon (1971), and have led to the transfer of this species to Scrippsiella. Material and Methods Isolate UTEX 1948 was grown in medium GPM (Loeblich, 1975) at 30 C over 500 ft-c continuous cool white fluorescent light. Cells were fixed for scanning electron microscopy as previously described (Loeblich & Sherley, 1979). The theca was stained and examined using the light microscope as previously described (Schmidt et al., 1978). Exponentially growing cells were measured at 400% after fixing with Lugol’s (I, KI) solution. 46 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Figs. 1-4. Scrippsiella gregaria. 1, Epithecae with 3a and 6” plates viewed from exterior; 2, Same, viewed from interior; 3 and 4, Epithecae with 4a and 7’ viewed from exterior; note wide sutures in Fig. 3. Scale bars = 10 um. Results DIVISION PYRRHOPHYTA PASCHER, 1914 ORDER PERIDINIALES HAECKEL, 1894 FAMILY PERIDINIACEAE EHRENBERG, 1832 Scrippsiella gregaria (LOMBARD & CAPON) COMB. NOV. Basionym: Peridinium gregarium Lombard & Capon, 1971:187, text-figs. 1LA-B, 2A-C. The most common thecal plate arrangement is: a pore plate, a preapical plate, 4 apical plates, 3 intercalary plates, 6 precingular plates, 6 cingular plates, 5 sulcal plates, 5 postcingular plates and 2 antapical plates (Figs. 1, 2). Four variations have been found in this tabulation: 1) 5—6 apicals, 2) 4— VOLUME 92, NUMBER 1 47 Figs. 5-8. Scrippsiella gregaria. 5 and 6, Cingular and sulcal plates indicated; 7 and 8, Sulcal and hypothecal plates labelled; note wide sutures in Fig. 7. Scale bars = 10 um. 6 intercalary plates (Figs. 3, 4), 3) 7 precingular plates, and 4) occasionally, a posterior intercalary plate present on the left side of the cell (Figs. 5, 6). Faint indications of vertical striations were noted on the dorsal epithecal plates (Fig. 3). Measurements of 30 cells of S. gregaria gave a mean length of 28.3 wm (range: 24.5—33.1 wm) and a mean width of 26.1 wm (range: 22.0- 31.8 wm). The ratio of mean length to mean width is 1.08. Discussion Comparison of the line drawing of Lombard & Capon (1971) to our mi- crographs suggests that the tabulation for S. gregaria given by Lombard & PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON VOLUME 92, NUMBER 1 49 Capon is in error. Specifically, in reference to their Fig. 1, only a small part of plate 6” is shown in ventral view and none of it in dorsal view. Their illustration of plate 6” is more consistent with it being part of their 3a plate. This reinterpretation decreases the number of precingulars to six which is the common number present in this series. In reference to their Fig. 1 the following comments are offered: 1) The plate 1’ ends at the anterior margin of the cingulum on the left side of the cell, while all cells of this species we have observed have the anterior sulcal extending into the epitheca where it meets the posterior margin of plate 1’. 2) On the dorsal side they figure three intercalary plates and show plate 3a as an isolated intercalary. A similar arrangement occurs in P. sociale ac- cording to Biecheler (1935). We have always found the intercalary plates to be in contact with each other and situated in such a position that there is never any contact of plate 3’ and 4”. Contact of these two plates was the basis for the section Sociale Biecheler, 1935, and Biecheler’s separation of P. sociale from other Peridinium species. 3) Our observations are in agree- ment with respect to the number of postcingular and antapical plates, but we have occasionally found a posterior intercalary on the left side of the hypotheca. This intercalary may be the same plate that Lombard & Capon illustrate as the 3’” in their Fig. 1A. 4) We find the sulcus made of five plates with the posterior sulcal plate the largest and the anterior sulcal extending into the epitheca. No pore was found in the sulcal region as illustrated by Lombard & Capon (their Fig. 1A, 2C). Because of the presence of a pore plate, preapical plate, six cingular plates and the marine habitation of S. gregaria, this species is removed from Peridinium and placed in Scrippsiella which is characterized by these fea- tures. Acknowledgments We thank E. Seling for SEM expertise and L. A. Loeblich for critical reading of the manuscript. This research was supported by the National Science Foundation (DEB 77-24620). Literature Cited Biecheler, B. 1952. Recherches sur les péridiniens.—Suppl. Bull. Biol. France Belgique 36:i- vi, 1-149. — Figs. 9-14. Scrippsiella gregaria. All at 10 kV except Fig. 2 at 30 kV. Scale bars = 10 wm. 9, Left ventral view; 10, Dorsal view; 11, Left-dorsal view of cell with 6 apical and 5 intercalary plates; 12, Dorsal view of cell with 6 intercalary plates; 13, Cells with posterior intercalary in posterior view; 14, Same, ventral view. 50 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Lee, R. F., and A. R. Loeblich, III. 1971. Distribution of 21:6 hydrocarbon and its relationship to 22:6 fatty acid in algae.—Phytochem. 10:593-602. Loeblich, A. R., HI. 1975. A seawater medium for dinoflagellates and the nutrition of Ca- chonina niei.—Jour. Phycol. 11:80-86. , and J. L. Sherley. 1979. Observations on the theca of the motile phase of freeliving and symbiotic isolates of Zooxanthella microadriatica (Freudenthal) comb. nov.—Jour. Mar. Biol. Assoc. U.K. 59:195—205, pls. 1-4. Lombard, E. H., and B. Capon. 1971. Peridinium gregarium, a new species of dinoflagel- late.—Jour. Phycol. 7:184—187. Schmidt, R. J., V. D. Gooch, A. R. Loeblich, HI, and J. W. Hastings. 1978. Comparative study of luminescent and nonluminescent strains of Gonyaulax excavata (Pyrrhophy- ta).—Jour. Phycol. 14:5-9. (ARL, III) Department of Biology, University of Houston, Houston, Tex- as 77004; (JLS) The Biological Laboratories, Harvard University, Cam- bridge, Massachusetts 02138; (RJS) Department of Botany, Duke Univer- sity, Durham, North Carolina 27706. PROC. BIOL. SOC. WASH. 92(1), 1979, pp. 51-69 POECILOSTOME COPEPODS (LICHOMOLGIDAE) FROM THE ALCYONACEAN CORAL CESPITULARIA MULTIPINNATA IN THE MOLUCCAS Arthur G. Humes and Masahiro Dojiri Abstract.—The alcyonacean coral Cespitularia multipinnata in the Mo- luccas is the host for three lichomolgid copepods. Paramolgus extendens, new species, is notable for its unusually long caudal rami (in the female with the ratio 16.8:1). Paramolgus pollicaris, new species, is distinctive in having a thumblike process on leg 5 in the female. Zamolgus cracens, new species, may be distinguished from its congeners by the long slender genital segment and the elongate free segment of leg 5 in the female (about as long as the genital segment). Forty-seven species of lichomolgid copepods are associated with Alcyo- nacea (Humes and Stock, 1973; Humes, 1973, 1975). Only one species has been reported from Cespitularia, namely, Zamolgus tridens Humes and Stock, 1973, from Cespitularia turgida Verseveldt in northwestern Mada- gascar. This paper contains descriptions of three new lichomolgids from Cespitularia multipinnata (Quoy and Gaimard) in the Moluccas. The alcyonaceans were isolated in plastic bags as soon as they were collected. Later they were gently washed in sea water containing about 5% ethyl alcohol. The water was then passed through a fine net and the cope- pods recovered from the sediment which was retained. When the alcyonaceans were collected, they were assumed to represent a single species, later identified by Dr. J. Verseveldt as Cespitularia mul- tipinnata. However, Dr. Verseveldt discovered a few colonies of a second species, Anthelia amboinensis (Burchardt), attached to the same fragments of dead coral. The small polyps and narrow flat stolons of this Anthelia seem to preclude it as the host for the many copepods recovered (about 435). Thus we consider Cespitularia multipinnata as the host of the cope- pods described below. The copepods were collected by the first author during the Alpha Helix East Asian Bioluminescence Expedition, which was supported by the Na- tional Science Foundation under grants OFS 74 01830 and OFS 74 02888 to the Scripps Institution of Oceanography and grant BMS 74 23242 to the University of California, Santa Barbara. The study of the copepods was aided by NSF grant DEB 77 11879. Sy PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Figs. 1-8. Paramolgus extendens, new species, female: 1. Dorsal (A); 2. Urosome, dorsal (B). 3. Genital area, dorsal (C). 4. Caudal ramus, dorsal (D). 5. Rostrum, ventral (E). 6. First antenna, with three dots indicating positions of aesthetes in the male, dorsal (D). 7. Second antenna, postero-inner (D). 8. Labrum, with paragnaths indicated by broken lines, ventral (F). VOLUME 92, NUMBER 1 53 We are greatly indebted to Dr. J. Verseveldt, Zwolle, The Netherlands, for the identification of the alcyonaceans. The observations and measurements were made on specimens cleared in lactic acid. All figures were drawn with the aid of a camera lucida. The letter after the explanation of each figure refers to the scale at which it was drawn. The abbreviations used are: A, = first antenna, A, = second antenna, L = labrum, MXPD = maxilliped, and P, = leg 1. Lichomolgidae Kossmann, 1877 Paramolgus Humes and Stock, 1972 Paramolgus extendens, new species Figs. 1—28 Type-material. —124 22, 141 36, and 106 copepodids from 15 colonies of the alcyonacean Cespitularia multipinnata (Quoy and Gaimard), in 5 m, southwestern side of Goenoeng Api, Banda Islands, 4°31'45”S, 129°51'55"E, 25 May 1975. Holotype 2 (USNM 171298), allotype (USNM 171299), and 255 paratypes (118 2 2, 137 63d) (USNM 171300) deposited in the National Museum of Natural History, Smithsonian Institution, Washington, D.C.; the remaining paratypes (dissected) and the copepodids in the collection of the first author. r Female .—Body (Fig. 1) moderately slender. Length (not including setae on caudal rami) 1.38 mm (1.27—1.44 mm) and greatest width 0.44 mm (0.39- 0.49 mm), based on 10 specimens in lactic acid. Ratio of length to width of prosome 1.55:1. Ratio of length of prosome to that of urosome 1:1. Segment bearing leg 5 (Fig. 2) 92 x 211 wm. Genital segment 194 x 194 pm, as long as wide, in dorsal view with slightly rounded lateral margins, broadest anteriorly and tapered a little posteriorly. Genital areas located dorsally near middle of segment. Each area (Fig. 3) with 2 small setae about 9 um long and a spiniform process. Three postgenital segments (Fig. 2) from anterior to posterior 70 x 113 wm, 59 x 97 wm, and 70 x 92 um. Anal Segment with posteroventral row of minute spinules on each side. Caudal ramus (Fig. 4) greatly elongate, 227 um long, 24 um in greatest width proximally, and 13.5 wm wide distally, ratio of length to distal width 16.8:1. Outer lateral seta 52 wm and naked, dorsal seta 23 wm and naked, outermost terminal seta 60 wm and innermost terminal seta 73 wm, both lightly feathered proximally. Two median terminal setae 122 um (outer) and 165 wm (inner), both smooth. Terminal flange with a few minute spinules. (Setal lengths approximate on account of being recurved or broken.) Body surface with small hairs (sensilla) as in Figs. 1 and 2. Egg sac approximately 384 x 184 um, containing about 14 eggs ranging from 81-108 um in diameter. (All egg sacs seen with eggs partly hatched or damaged.) 54 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON ZA = Lb Ms i, HY el Vi Aly ) \ ( AREA Y/ fi RIN [eH Be } AN A GIRSIN tN Ri RY ¥ AN RY wy Y AY) % Way ¥, REESE ZZ > PLL om LEE o LZ, Mee wa Le Wee LE SS TIAL 23 Figs. 9-17. Paramolgus extendens, new species, female: 9. Mandible, posterior (C); 10. | Paragnath, ventral (G); 11. First maxilla, anterior (C); 12. Second maxilla, posterior (F). 13. — Maxilliped, posterior (F). 14. Area between maxillipeds and first pair of legs, ventral (H). 15. Leg | and intercoxal plate, anterior (D). 16. Leg 2, anterior (D). 17. Third segment of endopod | of leg 3, anterior (D). VOLUME 92, NUMBER 1 5 Rostrum (Fig. 5) with rounded posteroventral margin. First antenna (Fig. 6) 310 um long. Lengths of 7 segments (measured along posterior nonsetiferous margins): 38 (59 wm along anterior margin), 84, 27, 43, 43, 32, and 22 um respectively. Formula for armature: 4, 13, 6, 3,4 + 1 aesthete, 2 + 1 aesthete, and 7 + | aesthete. All setae smooth. Second antenna (Fig. 7) 218 wm long including claw. Armature: 1, 1, 3, and claw + 5 setules. Fourth segment 57 um along outer side, 34 wm along inner side, and 25 wm wide. Claw 48 um along axis. Labrum (Fig. 8), mandible (Fig. 9), paragnath (Fig. 10), and first maxilla (Fig. 11) resembling in major respects those of Paramolgus politus (Humes and Ho, 1967). Second maxilla (Fig. 12) with proximal tooth on lash more spiniform than more distal teeth. Maxilliped (Fig. 13) with second segment bearing 2 unequal setae, shorter seta finely barbed, longer seta with a few hairs along one side. Small third segment with 2 barbed spines and a smooth seta. Ventral area between maxillipeds and first pair of legs (Fig. 14) slightly protuberant. Legs 1-4 (Figs. 15, 16, 17, 18) segmented and armed as in other Para- molgus. Barbs on outer side of terminal spine on third segment of exopod of leg 1 large and dentiform. Leg 4 with inner coxal seta 13 wm and naked. Exopod 156 um. First segment of endopod 44 um long xX 36 wm wide (with- out spiniform processes), its distal inner plumose seta 36 wm. Second seg- ment 70 < 31 um, its 2 unequal barbed spines 25 um (outer) and 52 um (inner). Both segments with outer marginal spinules rather than hairs. Leg 5 (Fig. 19) with elongate ventrally concave free segment 194 x 59 jum, in ovigerous specimens held around anterior part of egg sac. Two ter- minal setae 62 wm and 38 wm. Dorsal surface of free segment with small hairlike spinules. Between small dorsal seta and insertion of free segment a patch of spinules. Leg 6 represented by 2 setae and spiniform process on genital area (Fig. 3). Living specimens in transmitted light opaque, eye red, egg sacs gray. Male .—Body (Fig. 20) slender. Length (excluding setae on caudal rami) 1.14 mm (1.08—1.21 mm) and greatest width 0.31 mm (0.30—0.32 mm), based on 10 specimens in lactic acid. Ratio of length to width of prosome 1.71:1. Ratio of length of prosome to that of urosome 0.91:1, urosome slightly longer than prosome. Segment of leg 5 (Fig. 21) 38 x 65 wm. Genital segment 216 x 178 um, longer than wide. Four postgenital segments from anterior to posterior 35 x 62 wm, 32 x 59 wm, 24 x 59 um, and 35 x 57 wm. Caudal ramus similar to that of female but a little shorter, 208 x 14 um, ratio 14.9:1. Body surface with hairs (sensilla) as in female. 56 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Rostrum like that of female. First antenna similar to that of female but 3 aesthetes added (at points indicated by dots in Figure 6). Second antenna (Fig. 22) with inner spines on first and second segments. Labrum (Fig. 23) with both lobes having a posteriorly directed digitiform process and a minute bifurcate knob. Mandible, paragnath, first maxilla, and second maxilla like those of female. Maxilliped (Fig. 24) with second segment bearing 2 naked setae and 2 rows of spines. Claw 233 um along its axis, weakly divided midway, with a finely pectinate fringe along its concave margin and bearing 2 unequal proximal setae, larger seta bilaterally pectinate distally, smaller seta smooth. Ventral area between maxillipeds and first pair of legs as in female. Legs 1-4 segmented and armed as in female, except for third segment of endopod of leg 1 (Fig. 25) where formula is I, I, 4 instead of I, 5 as in female. Leg 5 (Fig. 26) with small unornamented free segment 25 x 11 um, ratio Depolel Leg 6 (Fig. 27) consisting of posteroventral flap on genital segment bearing 2 naked setae 33 wm and 26 um. Spermatophore 187 < 81 wm without neck, attached to female in pair (Fig. 28). Color in living specimens resembling that of female. Etymology .—The specific name extendens, from Latin extendo meaning to stretch out or prolong, alludes to the unusually long caudal rami in this species. Comparison with other species of Paramolgus.—Paramolgus extendens may be distinguished from all its congeners by the unusually long caudal rami, ratio in the female 16.8:1. In all other Paramolgus the ratio is less than 5:1. Paramolgus pollicaris, new species Figs. 29-57 Type-material.—14 22,5 3¢ from 15 colonies of the alcyonacean Ces- pitularia multipinnata (Quoy and Gaimard), in 5 m, southwestern side of Goenoeng Api, Banda Islands, 4°31'45”S, 129°51'55"E, 25 May 1975. Ho- lotype 2 (USNM 171301), allotype (USNM 171302), and 11 paratypes (9 22,2 36d) (USNM 171303) deposited in the National Museum of Natural History, Smithsonian Institution, Washington, D.C.; the remaining para- types (dissected) in the collection of the first author. Female .—Body (Fig. 29) with prosome moderately slender. Length (not including setae on caudal rami) 1.46 mm (1.33-1.53 mm) and greatest width 0.61 mm (0.56-0.63 mm), based on 10 specimens in lactic acid. Ratio of length to width of prosome 1.53:1. Ratio of length of prosome to that of urosome 1.71:1. VOLUME 92, NUMBER 1 57 = < SS <2 SSS — ose <> <2 SSS LF, ZS mz j i) { <> Ni \ Ny \ Figs. 18-19. Paramolgus extendens, new species, female: 18. Leg 4 and intercoxal plate, anterior (D). 19. Leg 5, dorsal (D). Figs. 20-28. Paramolgus extendens, new species, male: 20. Dorsal (A); 21. Urosome, dor- sal (E). 22. Second antenna, postero-inner (D). 23. Labrum, ventral (C); 24. Maxilliped, inner (1). 25. Endopod of leg 1, anterior (D). 26. Leg 5, dorsal (C). 27. Leg 6, ventral (D). 28. Spermatophores, pair attached to female, dorsal (J). 58 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Figs. 29-37. Paramolgus pollicaris, new species, female: 29. Dorsal (A); 30. Urosome, dorsal (E). 31. Segment of leg 5 and genital segment, lateral (K). 32. Genital area, dorsolateral (F). 33. Caudal ramus, dorsal (D). 34. Egg sac (B). 35. Rostrum, ventral (E). 36. First antenna, dorsal (I). 37. Second antenna, posterior (D). VOLUME 92, NUMBER 1 59 Segment of leg 5 (Fig. 30) 86 x 189 um. Genital segment elongate, 227 x 162 wm, in dorsal view with nearly parallel sides. Genital areas located laterally just anterior to midregion of segment (Figs. 30, 31). Each area (Fig. 32) with 2 minute setae about 7 wm long and a small blunt process. Three postgenital segments (Fig. 30) from anterior to posterior 70 < 86 um, 41 x 84 wm, and 51 xX 86 wm. Anal segment with smooth posteroventral margin. Caudal ramus (Fig. 33) short, 54 x 39 um, ratio 1.38:1. Outer lateral seta 110 wm and dorsal seta 35 wm, both smooth. Outermost terminal seta 165 ym, innermost terminal seta 229 wm, and 2 long median terminal setae 275 ym (outer) and 330 um (inner), both inserted between dorsal flange with minute marginal spinules and smooth ventral flange. All 4 terminal setae haired. Body surface with hairs (sensilla) and minute refractile points as in Fig- tne 29, Single egg sac observed (lacking 1 or 2 eggs) oval (Fig. 34), 483 x 276 wm, containing approximately 27 eggs with diameters ranging from 108- 119 um. Rostrum (Fig. 35) broadly linguiform. Elongate V-shaped sclerotization between bases of second antennae. First antenna (Fig. 36) 539 um long. Lengths of 7 segments (measured along posterior nonsetiferous margins): 49 (84 wm along anterior margin), 138, 38, 103, 81, 65, and 30 um respectively. Formula for armature: 4, 13 + 2 aesthetes, 6, 3 + 1 aesthete, 4 + 1 aesthete, 2 + 1 aesthete, and 7 + 1 aesthete. All setae smooth and relatively short. Second antenna (Fig. 37) 321 um long including claw. Armature as in Paramolgus extendens described above. Fourth segment 44 «wm along outer side, 25 wm along inner side, and 25 wm wide. Claw 43 um along its axis. Labrum (Fig. 38) with 2 broad posteroventral lobes. Mandible (Fig. 39) resembling in general that of P. extendens. Paragnath a small hairy lobe. First maxilla (Fig. 40) much like that of Zamolgus cracens described below. Second maxilla (Fig. 41) also similar to that of Z. cracens. Maxilliped (Fig. 42) with second segment bearing 2 very unequal setae, longer seta barbed, shorter seta smooth; segment with a few spinules on inner distal margin. Third segment (Fig. 43) with a unilaterally barbed spine and a small naked seta; segment terminating in a small smooth outer lobe and a blunt spiniform process bearing 9 small teeth. Ventral area between maxillipeds and first pair of legs (Fig. 44) slightly protuberant. Legs 1-4 (Figs. 45, 46, 47, 48) segmented and armed as in other Para- molgus. Leg 4 with inner coxal seta 35 wm and naked. Exopod 195 um. First segment of endopod 41 um long (without spiniform processes) x 40 um wide, its inner distal feathered seta 70 wm. Second segment 77 x 36 60 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON SS S = SEES rata = See YEE V KEP REA —. 25 L Figs. 38-46. Paramolgus pollicaris, new species, female: 38. Labrum, with paragnaths indicated by broken lines, ventral (F). 39. Mandible, posterior (F). 40. First maxilla, posterior (C). 41. Second maxilla, postero-inner (F). 42. Maxilliped, posterior (F). 43. Third segment of maxilliped, anterior (L). 44. Area between maxillipeds and first pair of legs, ventral (D). 45. Leg | and intercoxal plate, anterior (H). 46. Leg 2, anterior (H). 61 VOLUME 92, NUMBER 1 SSSR = SSSR ee SSS SOS SSS StS SOS < > SSS Z SES Waiters LORE FESO Zs SS LES ma Pe Ss Loe EFI LE LEE LZ yarace ZS ZS rae ze = TE LPez mwa Sa ue ease — = = = e = = = Se = = = = = Cette pes YQ Figs. 47-49. Paramolgus pollicaris, new species, female: 47. Endopod of leg 2, anterior (H). 48. Leg 4 and intercoxal plate, anterior (I). 49. Leg 5, dorsal (D). Figs. 50-57. Paramolgus pollicaris, new species, male: 50. Dorsal (A). 51. Urosome, dorsal (E). 52. Second antenna, posterior (D). 53. First and second segments of second antenna, ventral (M). 54. Maxilliped, inner (I). 55. Endopod of leg 1, anterior (D). 56. Leg 5, dorsal (C) 57. Leg 6, ventral (M). 62 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON ym, its 2 barbed terminal spines 51 wm (outer) and 76 wm (inner). Both segments haired along outer margins. Leg 5 (Fig. 49) with inner margin of unornamented free segment having a distally directed thumblike process. Dimensions: 59 um long, 30 wm wide proximally, and 38 wm wide at level of thumblike process. Two smooth terminal setae 57 wm and 43 wm. Leg 6 represented by 2 setae on genital area (Fig. 32). Living specimens in transmitted light opaque, eye red, egg sacs gray. Male .—Body (Fig. 50) a little more slender than in female. Length (with- out setae on caudal rami) 1.22 mm (1.20—1.26 mm) and greatest width 0.43 mm (0.42—0.43 mm), based on 4 specimens in lactic acid. Ratio of length to width of prosome 1.65:1. Ratio of length of prosome to that of urosome 1.3920" Segment of leg 5 (Fig. 51) 81 x 140 um. Genital segment elongate, 232 x 205 wm. Four postgenital segments from anterior to posterior 41 < 86 um, 46 x 78 um, 30 < 70 wm, and 46 x 73 um. Caudal ramus resembling that of female but smaller, 44 x 33 um, ratio 33316 Body surface with hairs (sensilla) and refractile points as in female. Rostrum and first antenna like those of female. Second antenna (Fig. 52) with first and second segments bearing raised ornaments (modified spines ?), each flattened and ribbed and resembling a scallop shell (Fig 53). Third segment with row of 4—5 spinules. Labrum, mandible, paragnath, first maxilla, and second maxilla like those in female. Maxilliped (Fig. 54) with second segment bearing 2 setae, one smooth, other pectinate along one side, and 2 rows of spines. Claw 288 wm along its axis, bearing proximally 2 very unequal setae, small seta smooth, large seta distally with closely spaced delicate hairs giving appearance of a striated membrane. Ventral area between maxillipeds and first pair of legs as in female. Legs 1—4 segmented and armed as in female, except for third segment of endopod of leg | (Fig. 55), where formula is I, I, 4 instead of I, 5 as in female. Leg 5 (Fig. 56) with small unornamented free segment 26 x 11 um, ratio 2. 30ul Leg 6 (Fig. 57) consisting of posteroventral flap on genital segment bearing 2 naked setae about 26 um long. Spermatophore not seen. Color in living specimens similar to that of female. Etymology.—The specific name pollicaris, Latin meaning of a thumb, alludes to the thumblike process on leg 5 in the female. Comparison with other species of Paramolgus.—Paramolgus pollicaris may be separated from all its congeners by two features: in the female by VOLUME 92, NUMBER 1 63 Figs. 58-68. Zamolgus cracens, new species, female: 58. Dorsal (A). 59. Urosome, dorsal (B). 60. Genital area, dorsal (F). 61. Caudal ramus, dorsal (D). 62. Rostrum, ventral (E). 63. First antenna, with three dots indicating positions of aesthetes in the male, dorsal (I). 64. Second antenna, posterior (I). 65. Labrum, with paragnaths indicated by broken lines, ventral (F). 66. Mandible, posterior (F). 67. First maxilla, posterior (C). 68. Second maxilla, posterior (F). 64 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON the thumblike process on leg 5, and in the male by the ornamentation re- sembling scallop shells on the first two segments of the second antenna. P. pollicaris is unusual in that the first antenna of the female has two aesthetes on the second segment and one on the fourth segment, thus pro- ducing the same formula as in the male. A similar condition occurs in My- coxynus villosus Humes (in press), a lichomolgid copepod from the fungiid coral Herpolitha limax (Esper). Zamolgus Humes and Stock, 1973 Zamolgus cracens, new species Figs. 58-83 Type-material.—27 22, 18 66 from 15 colonies of the alcyonacean Ces- pitularia multipinnata (Quoy and Gaimard), in 5 m, southwestern side of Goegoeng Api, Banda Islands, 4°31'45"S, 129°51'55"E, 25 May 1975. Ho- lotype 2 (USNM 171304), allotype (USNM 171305), and 36 paratypes (22 22,14 646) (USNM 171306) deposited in the National Museum of Natural History, Smithsonian Institution, Washington, D.C.; the remaining paratype (dissected) in the collection of the first author. Female .—Body (Fig. 58) with moderately slender prosome. Length (not including setae on caudal rami) 1.52 mm (1.46—1.63 mm) and greatest width 0.60 mm (0.56-0.63 mm), based on 10 specimens in lactic acid. Ratio of length to width of prosome 1.62:1. Ratio of length of prosome to that of urosome 1.58:1. Segment bearing leg 5 (Fig. 59) 108 x 216 wm. Genital segment elongate and slender, 221 x 162 um in greatest dimensions, slightly wider in anterior half than posteriorly. Genital areas situated laterally just in front of middle of segment. Each area (Fig. 60) with 2 small naked setae 11 wm and 7 wm and a small blunt process. Three postgenital segments (Fig. 59) from anterior to posterior 81 x 103 wm, 65 x 97 wm, and 70 x 108 wm. Anal segment with smooth posteroventral margin. Caudal ramus (Fig. 61) short, 59 x 49 um, a little longer than wide, ratio 1.22:1. Outer lateral seta 108 um, dorsal seta about 30 um, outermost ter- minal seta 194 um, innermost terminal seta 232 wm, and 2 median terminal setae 340 um (outer) and 454 um (inner), both inserted between small flanges, dorsal flange smooth, ventral flange with marginal row of minute spinules. All setae with lateral hairs except smooth dorsal seta. Body surface with hairs (sensilla) and refractile points as in Figure 58. Complete egg sac not seen, but individual eggs irregular in diameter, with average about 108 um. Rostrum (Fig. 62) linguiform. First antenna (Fig. 63) 510 wm long. Lengths of 7 segments (measured along their posterior nonsetiferous margins): 65 (89 wm along anterior mar- VOLUME 92, NUMBER 1 SSSss SS => =F S SS LLLEE See

*>......-... . 55 6 — Uropod 3 with outer ramus not exceeding 74 length of innerramus.. 7 6. Mandible palp article 1 distinctly shorter than palp article 3. Inner plate of maxilla 1 with 4 setae. Dorsal tooth of urosome segments i=? long: Coxa 4 broad’ ye o. cca ss cic oe eee See I. bowenae — Mandible palp article 1 as long as article 3. Inner plate of maxilla 1 with 2 setae. Dorsal tooth on urosome segments 1—2 short. Coxa 4 TRATION ie eties conse’! bs sdhat a « yeh Sy eaaee oro dey Mh a elak ORs tee Rea I. janisae 7. Palm of gnathopod 1 in males convex, entire ................ ie eis — Palm of gnathopod 1 in males with teeth or excavations .......... 9 8. Metasome segments 2-3 and urosomite 1 minutely serrate dorsally Berane ieee REM a oe yw Se aca olds cine 1 al eae I. curvidactyla — Metasome segments 1-3 and urosomites 1-2 minutely serrate dor- Seal pa rata tes tisaset ss tied ar + sdse.speagtiasme’s 6c 0 Re eR eee ee I. serra 9. Antenna | in males longer than antenna 2, with main flagellum longer ——————— ————— * SS VOLUME 92, NUMBER 1 83 than peduncle. Telson less than 1.5 times longer than broad ...... (OES OO Oe Oe ee Ge es ee ee ee I. chilkensis — Antenna 1 in males half as long as antenna 2, with main flagellum shorter than peduncle; telson 2.5 times longer than broad... I. nagatai Acknowledgments I am indebted to J. L. Barnard of Washington, D.C. and M. Bowen of Virginia for the help regarding the material and literature for realization of this work. Literature Cited Barnard, J. L. 1959. Liljeborgiid Amphipods of Southern California coastal bottoms, with a revision of the family.—Pacific Naturalist 1(4):12—28, figs. 1-12, 3 charts. Chevreux, E. 1920. Sur quelques amphipodes nouveaux ou peu connus provenant des cotes de Bretagne.—Bulletin de la Société zoologique de France 45:75-87. Chilton, C. 1921. Amphipoda. Fauna of the Chilka Lake.—Memoirs of the Indian Museum 5(8):519-558, 12 text figures. Imbach, M. C. 1967. Gammaridean Amphipoda from the South China Sea.—Naga Report 4:39-167, plates 1-33. Karaman, G. S. 1975. 65. Contribution to the knowledge of the Amphipoda. Two interesting amphipods from the Adriatic Sea, Iphimedia jugoslavica, n.sp. and Idunella pirata Krapp-Schickel 1975.—Poljoprivreda i Sumarstvo Titograd 21(3):31—48, figures 1-9. Krapp-Schickel, G. 1975. Neues tber die Liljeborgiiden des Mittelmeeres (Crustacea, Am- phipoda).—Bollettino del Museo Civico di Storia Naturale Verona I:455-—472, figures 1-8. Nagata, K. 1965. Studies on marine gammaridean Amphipoda of the Seto Inland Sea. I.— Publications of the Seto Marine Biological Laboratory 13(4):131-170, figs. 1-15. Sars, G. O. 1876. Prodromus descriptionis Crustaceorum ft [sic] Pycnogonidarum, qvae in expeditione Norvegica anno 1876, observavit.—Archiv for Mathematik og Naturviden- skab, 2:227-—271 [sic, really 337-271 in error]. . 1879. Crustacea et Pycnogonida nova in itinere 2do et 3tio expeditionis Norvegicae anno 1877 & 78 collecta. (Prodromus descriptionis).—Archiv for Mathematik og Na- turvidenskab 4:427-476. . 1895. Amphipoda. An account of the Crustacea of Norway with short descriptions and figures of all the species. Amphipoda. Universitetsforlaget Bergen and osla 1: viii and 1-711, 240 plates, 8 supplem. plates. Biological Institute, Titograd (Yugoslavia). PROC. BIOL. SOC. WASH. 92(1), 1979, pp. 84-97 ANATOMICAL NOTES ON LUTODRILUS MULTIVESICULATUS (ANNELIDA: OLIGOCHAETA)! Michael L. McMahan Abstract.—Anatomical studies of specimens of Lutodrilus multivesicu- latus McMahan, 1976, demonstrate the uniqueness of the species and of its monotypic family Lutodrilidae among the oligochaetes. Features conspic- uously different from those of other megadrile taxa include the arrangement of the parietal musculature, the large number of metameres containing male reproductive structures, and a vascular system that includes eleven pairs of hearts and caudal subcuticular capillary beds. The description of Lutodrilus multivesiculatus McMahan, 1976, placed this megadrile into a monotypic genus and family, the Lutodrilidae. Sub- sequent to the description, more extensive anatomical studies of the species have been completed. External Anatomy Secondary furrows are present anterior to xxi—xxiv, and tertiary furrows begin in vii, occurring posteriorly to xiv—xxiv. Each type of intrasegmental furrow is most evident dorsally. Body shape is almost circular in anterior cross section and is quadrangular posteriorly, beginning approximately at vil. The quadrangular shape is most pronounced posterior to gonadal seg- ments, with setal couples at the 4 angles of the body. Setal arrangement is lumbricin, with setae closely paired in a slightly postequatorial position. Setae are curved distally and have jagged, transverse ridges immediately proximal to the curved portion. A nodulus is one-third of the setal length from the distal tip. In nonregenerates, the tip of the tail often is reflected over the posterior border of the anus. Nephropores are inconspicuous, iden- tifiable only in histological sections. The clitellum is annular and only slightly swollen, with indistinct anterior and posterior margins. The body in this region is compressed dorsoven- trally, and the ventral surface is flattened or concave. The extent and lo- cation of the clitellum are variable, as indicated in Table 1. Intraclitellar ' This publication is derived from research conducted in partial fulfillment of requirements for the Ph.D. degree in the Department of Zoology and Physiology of Louisiana State Uni- versity, Baton Rouge. Appreciation is extended to Dr. Walter J. Harman for his guidance during this study. ne —geeeeeeeeeeeeeeeee VOLUME 92, NUMBER 1 85 Table 1. Location of clitellum in Lutodrilus multivesiculatus. Segments Number of Number of included segments specimens xxIxvi 47 | XXIxiv 45 | XxXIdxvi 46 | XxXIIXvili 48 | XxIxxl 5 | XXlItXv 44 1 XXlIIxv1 45 | XxV-Ixi 37) | intersegmental furrows are not obliterated by clitellar development. The clitellum is multilayered dorsal to the alae only, not extending onto the ventrum; however, the coloration is annular with the exception of the genital tumescences and ventral surfaces of the alae, both of which are white to tan in contrast to the reddened coloration of the clitellum. The tubercula pubertates include 16-32 segments, xxii-liii (Table 2), are aliform, and extend 1.5 mm from the body in preserved material and 2-3 mm in living animals (Fig. 1). The alae originate lateral to B and in life are flared laterally. Genital tumescences are small, rounded elevations not sharply demarcated from the surrounding epidermis (Fig. 1). Each tumesc- ence surrounds and includes 1 ventral setal pair, and the tumescences are superficial, being totally extramuscular. Each of segments xili-xxx, usually XVI—XXViil, and xxxiv—lii may possess 1, 2, or no tumescences, each limited to the region around a,b (Tables 3 and 4). Frequently, genital tumescences Table 2. Location of alae in Lutodrilus multivesiculatus. Segments Number of Number of included segments specimens XXlili 30 1 XXU-iii 32 1 XXVii-xlii 16 | XXVili—xlix Pep | xxviii 23 1 XXVili4i 24 2 XX1X—xlV1 18 1 XX1X—XIVili 20 1 XXIX—xlix 21 2 xxix! 22 3 XXIXii 2S 3 XXX—xlVi 17 2 XK DD, 1 86 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Figs. 1-2. Lutodrilus multivesiculatus. 1 (left), Anterior end of clitellate specimen of Lu- todrilus multivesiculatus, ventral view. a, ala; gt, genital tumescence; mt, male tumescence; op, location of oviducal pore; s, spermatophore; sp, location of spermiducal pore. 2 (right), Diagrammatic longitudinal section of genital region, not drawn to scale. 0, ovary; of, oviducal funnel; op, oviducal pore; s, spermatheca (spermathecae illustrated in only 3 intersegmentals); sfp, posteriormost spermiducal funnel; sp, spermiducal pore; sva, anteriormost seminal vesicle; Svp, posteriormost seminal vesicle; ta, anteriormost testis; tp, posteriormost testis; vd, vas deferens; ve, vas efferens. a a RR I I —— —-— — | | | VOLUME 92, NUMBER 1 87 Table 3. Distribution of genital tumescences in Lutodrilus multivesiculatus First and last segments of Number of Number of series segments specimens xiii 40 ] xvi 37 | XVi-xl vill 33) | Xvi-xlix 34 D xviii 36 2 XVi-lli 37 2 XVli-li 35 2 XViiiii 36 1 XVilii 34 1 xix 33 1 are absent on most or all of segments xxxiv—xlvi. In clitellate individuals, genital tumescences are white to tan in contrast to clitellar coloration where- as, in aclitellates, the color of the genital tumescences is like that of the surrounding epidermis. The male tumescence is an elevated, flattened area on the ventrum of xxxii—xxxill, sometimes extending onto portions of xxxi and/or xxxiv (Fig. 1). The male tumescence contacts the alae, thus including both ventral setal couples of each segment. The ventral setal pairs of xxxi- XXXIV are displaced medially. In xxxi and xxxiv, the amount of displacement is equal to half of AB whereas b of xxxii and xxxiii is in the A line of segments other than xxxi-xxxiv (Fig. 1). Spermathecal pores are minute, unrecognizable macroscopically, and are multiple in each intersegmental 15/ 16—25/26, ventral to C. Internal Anatomy Musculature Septa are present posteriorly beginning with 3/4 and are thickened and heavily muscularized 7/8—23/24. Posterior to the pharynx, septa are mark- edly depressed posteriad. Muscles extend from the pharynx to parietal in- Sertions, ending in vii. Anteriorly, the body wall is distinctive, being ca. 1.5 mm thick in ix of a mature specimen and having 5 distinct muscle layers: subepithelial circular, diffuse longitudinal, median circular, trabeculate longitudinal, and retroper- itoneal circular (Fig. 3). The median circular muscle layer diminishes pos- teriorly and ends in xviii. In midbody, the body wall is ca. 0.75 mm thick, and there are 3 muscle layers: 2 circular layers with a median trabeculate longitudinal layer (Fig. 4). The retroperitoneal circular layer is much thinner than anteriorly. Near the periproct, the body wall musculature is much 88 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Table 4. Pattern of distribution of genital tumescences of 3 specimens of Lutodrilus multi- vesiculatus. Position on segment Segment 1 D) 3 pre-xv none none none XV right none none XVI left bilateral none XVI bilateral left none XVill bilateral right left XIX bilateral right none OX bilateral left right XXl bilateral bilateral bilateral XXil bilateral bilateral right XXlil bilateral bilateral right XXIV bilateral bilateral bilateral XXV bilateral bilateral bilateral XXVI bilateral bilateral bilateral XXVil bilateral none bilateral XXVIII bilateral none left XX1X right none none XXX bilateral none none XXXI bilateral none right XXXII male tumescence male tumescence male tumescence XXXIil male tumescence male tumescence male tumescence XXXIV bilateral none none XXXV none none none XXXVI none none . none XXXVIl right none none XXXVIil left none none XXXIX none right none xl none none none xli left none none xlil none none none xliil none none none xliv none none none xlv none none none xlvi none none none xl vil bilateral bilateral none xl vill bilateral bilateral right xlix right right none l right none none li right none bilateral lii and following none none none reduced. The only distinct layer is the outer circular muscle, beneath which is a loosely arranged layer of longitudinal fibers that are not organized into well-defined bundles. Immediately internal to the peritoneum, there is a muscle layer consisting of | to 2 thicknesses of circular muscle fibers. | | : | VOLUME 92, NUMBER 1 89 In megadriles, the existence of distinct circular muscle layers other than immediately subepidermal in position is unknown. Such a condition would be expected to accompany reduction of the outermost layer of circular mus- cle; however, this is not the case. To the contrary, in L. multivesiculatus, the outermost circular muscle layer is thickest anterior to and including x— the part of the body in which the 2 inner circular layers are best developed. The proportional amount of circular muscle in this region is greater than in any other known megadrile and, comparatively, results in an extremely powerful probing capability with the anterior segments. The retroperitoneal circular layer, always present throughout the body, is thickest anterior to x, posterior to which it diminishes to equal the thick- ness of the outermost circular layer. Near the periproct, the retroperitoneal circular musculature is one-sixth of the thickness of the subepidermal layer. The trabeculate muscle layer is most highly developed in the region of Xil, where it is ca. 1.2 mm thick. ‘‘Kastchen”’ are poorly defined, and they lack discernible internal organization. Fibers within a ‘‘kastchen”’ are ar- ranged into small, randomly distributed bundles that are most dense ectally. Fascicles of connective tissue occasionally infiltrate between the bundles. This arrangement resembles that reported in Eisenia foetida (Savigny) and Allolobophora chlorotica (Savigny) by Harman (1960) although it is less organized than in either of these 2 species. The outer longitudinal muscle layer consists of small, diffuse bundles of fibers that only occasionally are enclosed by connective tissue fascicles. The radial fascicles of connective tissue separating ““kastchen’’ of the tra- beculate layer sometimes penetrate the median circular layer and pass through the outer longitudinal layer, dividing its fibers into ill-defined bun- dles. Nervous System The brain, located in ii—iii, is bilobed and gives rise to 2 circumpharyngeal commissures at 2/3. Each commissure is one-third the diameter of the single ventral nerve cord, which the commissures join in ill. Five pairs of nerves branch anteriad from the brain: 2 to the roof of the buccal cavity and 3 to the prostomium, peristomium, and parieties. Segmentally, 3 pairs of lateral nerve trunks arise from the nerve cord. Digestive System The digestive tube lacks crop, gizzard, and calciferous glands. The phar- ynx is in ii—vi, with muscle fibers to the parieties extending to vii. An esophageal valve originates from the dorsal wall of the esophagus in vii. The esophageal mucosa becomes folded longitudinally in xvi, with vascular sinuses in xvi-xx. The gut widens at 20/21, and intestinal pouches begin in 90 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Figs. 3-4. Lutodrilus multivesiculatus. 3 (left), Cross section of anterior body wall; 4 (right), Cross section of body wall in clitellar segment. a, subepithelial circular muscle; b, diffuse longitudinal muscle; c, median circular muscle; d, trabeculate longitudinal muscle; e, retro- peritoneal circular muscle; f, lateroparietal vessel. Scales 100 um. XX1-xxiV, extending to the region of 1. Each pouch, a dilatation of the intestine, is variable in length, extending up to 10 segments. The dorsal typhlosole begins as a low proliferation of tissue in xxxi—xxxiv and may obliterate the intestinal lumen posteriorly. The peduncle of the typhlosole is nearly as broad as the distal portion (Fig. 5). Chloragogen is present from the anterior segments, with greatest accumulations in xx and succeeding segments. Nephridia Nephridial tubules extend into the upper part of the coelom and adhere closely to the parieties. Nephrostomes are single. Genitalia Seminal vesicles are largest in xiv—xxii, where they often fill the coelom to the middorsal line (Fig. 2). Vesicles are attached to the posterior faces VOLUME 92, NUMBER 1 91 Fig. 5. Cross section of typhlosolar intestine of Lutodrilus multivesiculatus, scale 100 um. of their respective septa with the exception of the vesicles of xi and xii, which attach to the anterior faces of septa 11/12 and 12/13, respectively. Testes are digitate, ending distally in numerous strings. The large, polypli- cate spermiducal funnels surround ventrolateral portions of the seminal ves- icles. The vasa efferentia are microscopic, having diameters of 50-75 um. They enter the parieties one segment posterior to their respective funnels and orient posteriorly in a retroperitoneal position. One, or sometimes 2, segments posterior to their origins, the vasa efferentia join the intramuscular vasa deferentia, which also are microscopic (ca. 75 «~m in diameter). The vasa deferentia are embedded in the trabeculate longitudinal muscle layer approximately half the distance through the body wall; however, they move toward the coelom at the junction with each vas efferens. Each vas deferens 92 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON is lateral to B until 31/32, where it begins medial orientation through the ventral parieties toward the spermiducal pore in xxxil. Prostates are lacking. Each mature ovary resembles a rounded triangle with a posterior apex tapering to a single egg string containing 6-10 maturing oocytes that are visible through the ovarian wall (Gates, 1976; McMahan, 1976). Immature ovaries lack the egg string although developing oocytes are visible through the wall of the ovary (Gates, 1976). The crenellate oviducal funnels are smaller than and more closely attached to the septum than are the spermi- ducal funnels. Oviducts pass posteriorly through septum 23/24, which is depressed posteriad by the seminal vesicles of xxii, and the ducts then orient anteriad toward their entrance into the parieties lateral to B and presetal. Spermathecae are ovoidal, averaging ca. 150 x 200 82220 18%, 2S, ee PMR An SERA Cephalogammarus ieadordinary 0 20 eS 2 AS? ee Oe on 34 . Coxae 1-4 and article 2 of pereopods 5—7 strongly setose on mar- SUMS eee Te Se oe hata ee ee a ee ee Turcogammarus Coxae |-4 and article 2 of pereopods 5—7 with short and sparse STE i Na Rl NA BE i air dacstr cnatia dpe Dikerogammarus . Article 4 of pereopod 4 with only 4—5 posterior setae, (antenna 1 oPMiKerosummarus form)’. ee oe eee Akerogammarus Article 4 of pereopod 4 with 10+ posterior setae, (antenna 1 of LOnvOeUmmaras TOLM)') 2. oR ee 36 | { | | VOLUME 92, NUMBER 1 131 36. Article 2 on outer ramus of uropod 3 ‘‘elongate’’ (15+ percent of article 1), first 2 flagellar articles of antenna 2 together more than 95 percent as long as article 5 of peduncle ......... Stenogammarus — Article 2 on outer ramus of uropod 3 not elongate (less than 12 percent of article 1), first 2 flagellar articles of antenna 2 together less than 80 percent as long as article 5 of peduncle ............. a7 37. Posterior setae on article 4 of pereopod 4 divided into groups . mun reints | ee BR irs cor meta BPs yk epee wera Obesogammarus — Posterior setae on article 4 of pereopod 4 in continous fan........ 38 38. Inner ramus of uropod 3 about half as long as outer ramus ...... Re ee ORC ae MIGMOIIe sore tte leo) leit reys eel) Be Euxinia — Inner ramus of uropod 3 about one third as long as outer ramus... Pee eee eer AEs fer Kcwel, Gaga: Pontogammarus Baku, Karaman and Barnard, new genus Type-species.—Pontogammarus paradoxus Derzhavin, in Derzhavin and Pjatakova (1967). Named for the city on the Caspian Sea. Masculine. Body somewhat slender, urosomites free, at least segment 3 with 2 dorsal spines, others apparently flat and naked. Rostrum obsolescent, lateral ce- phalic lobes subquadrate, sinus present. Eyes very small. Antennae short, extending subequally; antenna 1 of Pontogammarus form (almost of Niphargoides form), no articles humped, ratio of peduncular articles = 26:10:6, ratio of flagella = 24:10, ventral setae of article 1 on peduncle weak and terminal, accessory flagellum 3-articulate. Ratio of ar- ticles 3,4,5 and flagellum of antenna 2 = 8:11:13:26 (to some extent of Steno- gammarus form), article’ 3 weakly humped. Mouthparts [unknown]. Coxae of medium size, setose, coxa | weakly curved anteriorly, coxa 4 lobed. Gnathopods 1-2 strongly subchelate, of medium size, gnathopod 1 enlarged but otherwise almost like gnathopod 2, wrists short (but shorter on gnathopod 1), scarcely lobed, hands weakly elongate, subrectangular, pal- mar slopes almost identical, oblique, short. Pereopods 3-7 moderately fossorial; article 4 of pereopod 4 narrow but with 10+ posterior groups of (very few) setae (each), article 5 weakly ex- panded, moderately setose posteriorly. Article 2 of pereopods 5-7 densely setose posteriorly, of pereopod 5 weakly expanded and scarcely lobate, of pereopod 6 unexpanded, tapering, unlobate, posteriorly excavate; of per- eopod 7 expanded, lobate. Pleopods ordinary. Epimeron 3 lacking posteroventral fan of setae, pos- teroventral corner with sharp, weakly curved tooth. Rami of uropods 1-2 extending equally, [peduncular and ramal spination or setation unknown]. Uropod 3 not extended beyond uropods 1-2, parviramous, outer ramus 132 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON short, weakly spinose and setose, article 2 short. Telson of ordinary length, lobes tapering, moderately setospinose apically. Gills [?2-6], broadly ovate, stalked. Oostegites of medium expansion. Relationship.—Differing from ordinary pontogammarid genera in en- larged gnathopod 1; from Iphigenella Sars in lack of prehensility on per- eopods 3-7; from Lanceogammarus new genus in the enlarged female gnathopod 2, which, though not as large as gnathopod 1, is significantly enlarged; from Pachyschesis Bazikalova, in curved coxa 1, lobed coxa 4, presence of long fully ventral setae on coxae 1-4, the distinctly lobate per- eopod 7 and distinctly though weakly enlarged gnathopod 1; from Amath- illina Sars in the uncarinate body and more strongly setose coxae and per- eopods; differing from Yogmelina brachyura by the 3-articulate accessory flagellum, by the sharp tooth on epimeron 3 and by the distinctly lobate basis of pereopod 7. Lobogammarus has gnathopod 1 smaller than gnath- opod 2. Species.—paradoxus (Derzhavin, in Derzhavin and Pjatakova, 1967); Caspian Sea, moderately fossorial, 1. Cephalogammarus Karaman and Barnard, new genus Type-species.—Gammarus macrocephalus Sars, 1896. Name.—Reference to encephalized Gammarus. Masculine. Body ordinary, urosomites free, 1-2 each with large elevated knob, all weakly spinose. Head greatly enlarged, about as long as first 2.5 pereonites, rostrum small, lateral cephalic lobes subquadrate, sinus present. Eyes pres- ent. Antennae elongate, antenna 1 slightly the longer, of Dikerogammarus form, no articles humped, ratio of peduncular articles = 30:21:12, ratio of flagella = 114:21, ventral setae of peduncular article 1 weak and terminal; accessory flagellum with 3-6 articles. Ratio of articles 3,4,5 and flagellum of antenna 2 = 12:22:20:34, article 3 unhumped. Mouthparts unknown, though unexpanded view of mandibular palp show- ing D and E setae, presumed to be as in Dikerogammarus. Coxae elongate, long setae absent, coxa 4 lobate. Gnathopods 1-2 strong- ly subchelate, large to medium, gnathopod 2 enlarged but like gnathopod 1, wrists short, weakly to poorly lobed respectively, hands elongate, palmar slopes almost identical, oblique, of medium length; female [unknown]. Pereopods 3-7 moderately fossorial; article 4 of pereopod 4 narrow, with 4-5 widely separated groups of sparse posterior setae, article 5 narrow, rectangular, with only 3-4 posterior setal groups. Article 2 of pereopods 5— 7 with short posterior setules, of pereopods 5—6 weakly expanded, unlobate, posterior margins both weakly concave (sinuous), of pereopod 7 expanded, lobate. | ' / VOLUME 92, NUMBER 1 153 Pleopods ordinary. Epimeron 3 lacking posteroventral fan of setae. Outer ramus of uropod 2 slightly shortened and lacking marginal spines, peduncle of uropods 1-2 apparently without setae. Uropod 3 extended, parviramous, outer ramus elongate, setose, weakly spinose, article 2 short. Telson of ordinary length (slightly short), almost fully cleft, apices of medium width, moderately setospinose. Gills [?2-6]. Oostegites [?slender]. Relationship.—Like Dikerogammarus but head immense. Species.—macrocephalus (Sars, 1896); Caspian Sea, moderately fosso- Half. Kuzmelina, Karaman and Barnard, new genus Name.—Contrived. Feminine. Type-species.—Gmelina kusnezowi Sowinsky, 1894. Body carinate, with bilateral dorsal lines of humps anterior to urosome, best developed on pereonite 6 to pleonite 3; pereonal pleurae humped. Uro- somites free, poorly armed. Rostrum short, lateral cephalic lobes protuber- ant, mammilliform, sinus obsolescent but rudimentary lateral wing tooth present (reminiscent of Axelboeckia). Eyes present, very close to anterior cephalic margin. _ Antennae of medium size and extension, almost of equal extent, antenna 1 slender, of Dikerogammarus form, ratio of peduncular articles = 30:24:15, flagellar ratio = 66:4, ventral setae of peduncular article 1 weak and mostly terminal, accessory flagellum I-articulate. Ratio of articles 3,4,5 and flagellum of antenna 2 = 15:33:32:42, no articles humped. Mouthparts unknown, space below for addition when described. Labrum [?broader than long, entire, rounded]. Mandibular incisor [?toothed, molar triturative, ratio of palp articles = 00:00:00, article 3 weakly falcate, setae = ABCDE]. Inner lobes of labium [?absent, weakly gaping]. Maxillae [?well setose medially, inner plate of maxilla 1 triangular, fully setose medially, outer plate with ?7 spines, palps asymmetric]. Inner plate of maxilla 2 [? with oblique facial row of setae]. Outer plate of maxilliped [?medially spinose, article 3 of palp unlobed, dactyl shorter than 3, unguiform, with nail]. Coxae long, setae short to medium, coxa 1 slightly curved forward, coxa 4 lobed, coxa 5 shorter than 4. Gnathopods medium to small, subchelate, male gnathopods of medium size, alike, wrists short-medium, scarcely lobed, hands elongate, palms oblique, excavate, well defined, of medium length; female gnathopods small, dissimilar, hands similar, rectangular, : palms oblique, short, wrists poorly lobed, wrist of gnathopod 1 short, of gnathopod 2 elongate or medium. Pereopods 3-7 not fossorial. Article 2 of pereopods 5-6 scarcely expanded and tapering, unlobate, of pereopod 7 weakly expanded, unlobate, of S—7 _ weakly setose posteriorly. 134 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Pleopods ordinary. Epimeron 3 lacking posteroventral fan of setae. Rami of uropods 1-2 extending equally, lacking marginal spines, peduncles poorly armed. Uropod 3 weakly extended, parviramous, outer ramus elongate, weakly spinose and setose, article 2 short. Telson short, broad, deeply cleft, apices strongly spinose, with lateral spines. Coxal gills [?2-6]. Oostegites [?slender]. Relationship.—Differing from Gmelina Sars in the doubled carina, the protruding cephalic lobes, the rudimentary cephalic wing, the marginal eyes and short telson. See Gmelinopsis. Species.—kusnezowi (Sowinsky, 1894) (Sars, 1894); Caspian and Azov seas, l. Lanceogammarus Karaman and Barnard, new genus Type-species.—Gammarus andrussowi Sars, 1896 (here selected). Name.—Lanceo—referring to thin gnathopod 2 of a Gammarus-like tax- on. Masculine. Body ordinary, urosomites free, unhumped, spinose. Rostrum obsoles- cent, lateral cephalic lobes subquadrate, sinus present. Eyes present. Antennae elongate, antenna | much longer than antenna 2, of Dikerogam- marus form, no articles humped, ratio of peduncular articles = 30:22:10, ratio of flagella = 150:18, ventral setae of peduncular article 1 weak and terminal, accessory flagellum multiarticulate. Ratio of articles 3,4,5 and fla- gellum of antenna 2 = 15:36:36:57, article 3 weakly humped. Labrum [?broader than long, entire, rounded]. Mandibular [?incisor toothed, molar triturative], ratio of palp articles = 4:13:10, article 3 weakly falcate, setae = BDE. Inner plate of maxilla 1 triangular, fully setose me- dially, outer plate with 10 (?11) serrate spines, palps [?asymmetric]. [?Inner plate of maxilla 2 with oblique facial row of setae]. Outer plate of maxilliped medially setose, article 3 of palp unlobed, dactyl shorter than 3, unguiform, [?with nail or not, illustration in Carausu, 1943, not showing nail]. Coxae of medium size, long setae absent, coxa 4 lobed. Gnathopods 1-2 strongly subchelate, diverse, medium to small, gnathopod 1 enlarged, with short scarcely lobed wrist, hand elongate, palm oblique, of medium length; gnathopod 2 small, thin, wrist elongate, scarcely lobed, hand almost as long as wrist, rectangular, palm oblique, very short; male gnathopod 1 larger than in female. Pereopods 3-7 weakly fossorial; article 4 of pereopod 4 narrow, with 2 widely separated groups of setae and spines, article 5 narrow, rectangular, with only 2 posterior setae-spine groups. Article 2 of pereopods 5-7 mod- erately expanded and moderately setose posteriorly, pereopods 5-6 scarcely to weakly lobate, pereopod 7 strongly lobate, article 2 of pereopod 6 convex posteriorly. | VOLUME 92, NUMBER | 135 Pleopods ordinary. Epimeron 3 lacking posteroventral fan of setae. Outer ramus of uropod 2 slightly shortened, lacking marginal spines, peduncle of uropod | with basofacial spine, no setae. Uropod 3 extended, parviramous, outer ramus elongate, weakly spinose, article 2 of medium length. Telson short, deeply cleft, apices tapering to medium width, spinose, also with basodorsal spine pairs. Gills [?2—6]. Oostegites slender. Relationship.—Differing from Akerogammarus, Dikerogammarus, Lo- bogammarus, Shablogammarus, and ‘‘Stenogammarus’’ macrurus of Car- ausu (1943) in the enlarged gnathopod |. See Baku; Iphigenella andrussowi formerly assigned to Iphigenella but differing from that genus in the non- prehensile pereopods. Species.—andrussowi (Sars, 1896); Caspian and Black seas and their trib- utaries, |. Stenogammarus Martynov Stenogammarus Martynov, 1924:41; Stock, 1974:85 (Gammarus macrurus Sars, 1894a, selected by Stock, 1974). Wolgagammarus Stock 1974:85 (valid subgenus, see below). Body slender, urosomites free, unhumped, scarcely setulate or weakly spinose. Rostrum small, lateral cephalic lobe rounded or subquadrate, sinus present. Eyes present. Antennae short to medium, extending subequally; antenna 1 of Ponto- gammarus form, no articles humped, ratio of peduncular articles = 35:13:8, flagellar formula = 58:15, ventral setae of peduncular article 1 weak and terminal, accessory flagellum multiarticulate. Ratio of articles 3,4,5 and fla- gellum of antenna 2 = 18:21:19:50, article 3 weakly humped, [antenna 2 of Stenogammarus form in which first 2 articles of flagellum together are al- most as long as peduncular article 5, and articles 4-5 of peduncle not humped, slender, article 5 the thinner, hence article 5 forming blend between article 4 and flagellum; this concept so variable and difficult to treat we abandon it as taxonomic device]. Mouthparts of type-species unknown but probably as follows: [Labrum broader than long, entire, rounded truncate. Mandibular incisor toothed, molar triturative, ratio of palp articles = 5:15:12, article 3 curved-linear, setae = AB(D)E, D and E setae terminal. Labium without inner lobes. Max- illae medially setose, inner plate triangular, fully setose medially, outer plate with 11 spines, palps ?asymmetric. Inner plate of maxilla 2 with oblique facial row of setae. Outer plate of maxilliped medially spinose, article 3 of palp unlobed, dactyl shorter than 3, unquiform, ?with nail]. Coxae of medium size to long, setae short and sparse, coxa 4 lobed. 136 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Gnathopods strongly subchelate, medium to small, gnathopod 2 of male slightly larger than but like gnathopod 1, wrists short, poorly lobate, hands elongate to stout, palmar slopes identical, oblique, of medium length; female gnathopods both much smaller than in male, almost identical in size, wrists much longer but wrist of gnathopod 2 much longer than in gnathopod 1, hands subrectangular, palms short. . Pereopods 3-7 moderately fossorial; article 4 of pereopod 4 with posterior setae arranged in bundles (type) or continuously (similis), article 5 narrow and rectangular (type) or expanded and trapezoidal (similis). Article 2 of pereopods 5-7 weakly (type) to strongly (similis) setose, in type article 2 of pereopod 5 not posteriorly setose, of pereopods 6—7 with short sparse setae, of pereopod 6 weakly expanded, posterior margin convex, of pereopods 5 and 7 strongly expanded, lobate. Pleopods ordinary. Epimeron 3 lacking posteroventral fan of setae. Rami of uropods 1-2 extending evenly, [?apparently outer ramus of uropod 2 lacking marginal spines, basofacial setae of uropod | apparently present]. Uropod 3 strongly extended, parviramous, peduncle slightly elongate (or not), outer ramus elongate, (spinose) and setose, article 2 elongate. Telson of ordinary length, apices of medium width, moderately spinose. Gills [?2-6]. Oostegites of medium breadth. Relationship.—Differing from Pontogammarus Sowinsky, Euxinia Tuc- olesco and Obesogammarus Stock in the elongate article 2 on the outer ramus of uropod 3. Notes.—Antennal measurements taken from carausui (=similis of Car- ausu, 1943); S. macrurus of Carausu (1943) removed to incertae sedis below. Variants.—Species other than the type with concave or sinuous posterior margin on article 2 of pereopod 6. Species.—carausui Derzhavin and Pjatakova, 1962 (=similis of Carausu, 1943, =olearii Derzhavin, 1951, =compresso-similis Carausu et et alia 1955 [nomen nudum], =karauschi Dedui, 1967, =kereuschui Mordukhai-Boltov- skoi and Ljakhov, 1972), compressus (Sars, 1894a), deminutus (Stebbing, 1906) (=minutus Sars, 1894a), macrurus (Sars, 1894a), similis (Sars, 1894a); Caspian, Azov and Black seas, attendant rivers, fossorial, 5 (see Steno- gammarus sp. below). Stenogammarus sp. Stenogammarus macrurus.—Carausu, 1943:59, plates 16, 17 (not Sars, 1894a). Like Stenogammarus but article 2 of pereopod 6 lobate; like Shablogam- marus Carausu, Dobreanu and Manolache, but article 2 on outer ramus of uropod 3 elongate, coxae 1-4 with long setae, antenna | of Pontogammarus form. VOLUME 92, NUMBER 1 137 Carausu’s material needs reexamination to see if it is properly described and, indeed, whether or not Sars’s depiction of S. macrurus is correct. If not, all species of Stenogammarus must be realigned to reflect the outcome. We do not know if Stock (1974) based some of the diagnosis of Stenogam- marus on this depiction by Carausu, a potential problem needing clarifica- tion. Species.—sp. (macrurus of Carausu, 1943); Danube River, fossorial, 1. Stenogammarus (Wolgagammarus) Stock S. (Wolgagammarus) Stock, 1974:85 (Stenogammarus dzjubani Mordukhai- Boltovskoi and Ljakhov, 1972, original designation). Like Stenogammarus but male gnathopod 2 of neotenic form, thus like female; article 5 of pereopods 3-4 especially elongate. Outer margin of uro- pod 3 with pinnate setae. Species.—dzjubani Mordukhai-Boltovskoi and Ljakhov, 1972; Volga Riv- er, man-made lakes, 1. Turcogammarus Karaman and Barnard, new genus Type-species.—Obesogammarus turcarum Stock, 1974. Name.—Type-species Turkish. Masculine. Body ordinary, or carinate, urosomites free, at least 1 and 2 with large elevated process (knob or tubercle), all spinose. Rostrum short, lateral ce- phalic lobes subquadrate, sinus present. Eyes present. Antennae of medium length, extending subequally, antenna 1 of Ponto- gammarus form, no articles humped, ratio of peduncular articles = 30:15:7, flagellar formula = 63:17, ventral setae of peduncular article 1 weak and terminal, accessory flagellum 4-articulate. Ratio of articles 3,4,5 and flagel- lum of antenna 2 = 17:27:21:50, article 3 weakly humped. Mouthparts of type-species only partly known, bracketed remarks need confirmation: labrum [?broader than long, entire, rounded]. Mandibular in- cisor [?toothed, molar triturative], ratio of palp articles = 5:12:9, article 3 weakly falcate, setae = ABCDE! Labium with inner lobes [?weak]. Max- illae [?medially setose, inner plate of maxilla 1 triangular, fully setose me- dially], outer plate with 9 spines, palps asymmetric. Inner plate of maxilla 2 with [?oblique facial row of setae]. Outer plate of maxilliped [?medially spinose, article 3 of palp unlobed, dactyl as long as 3, unguiform, with nail]. Coxae of ordinary length, strongly setose ventrally, coxa 1 weakly dilated distally, coxa 4 lobed. Gnathopods 1-2 subchelate, almost alike in shape, wrists short, scarcely lobed, hands ovate, palms oblique, of medium length, almost identical in slope, each with midpalmar spine, gnathopod 2 larger than 1. Female gnathopods smaller than in male. 138 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Pereopods 3-7 strongly fossorial; article 4 of pereopod 4 weakly expand- ed, with about 9 posterior bundles of setae, article 5 scarcely expanded, almost linear, strongly setose. Article 2 of pereopods 5-7 strongly setose posteriorly, moderately setose mediofacially, of pereopods 5 and 7 expand- ed, weakly and strongly lobed posteroventrally respectively, of pereopod 6 tapering and unlobed. Pleopods ordinary. Epimeron 3 lacking posteroventral fan of setae. Outer ramus of uropod 2 shortened, outer rami of uropods 1-2 without marginal spines. Uropod 3 [?weakly extended], almost parviramous, inner ramus short and scale-like but with several medial setae (by definition thus vari- ramous), outer ramus elongate, weakly spinose and strongly setose, article 2 short. Telson of ordinary length, cleft to base, apices tapering, with several spines and setae. Coxal gills [?2-6], [?o0void in type-species, ovoid in spandli]. Oostegites narrow. Variants.—Metasome with dorsomedial keel (spandli); gnathopodal palm slopes dissimilar (spandli). Relationship.—Differing from Dikerogammarus in the strong setosity of coxae, articles 4—5 of pereopods 34 and article 2 of pereopods 5-7; differing from Obesogammarus in presence of urosomal knobs. Species.—spandli (S. Karaman, 1931), turcarum (Stock, 1974); Northern Greece, springs and streams; Turkey, Mt. Ararat region, fountain (we place in Caucasus province), fossorial but ecology anomalous, 2. Yogmelina Karaman and Barnard, new genus Type-species.—Gmelina pusilla Sars, 1896. Name.—Contrived. Feminine. Body smooth, urosomites free, smooth or spinose. Rostrum short, lateral cephalic lobes protuberant, subacute to rounded, sinus obsolescent. Eyes present. Antennae of medium size and extending subequally; antenna | of Diker- ogammarus form, no articles humped, but peduncle stout, ratio of pedun- cular articles = 27:24:15, flagellar formula = 75:5, ventral setae of pedun- cular article | weak and terminal, accessory flagellum 1-articulate. Ratio of articles 3,4,5 and flagellum of antenna 2 = 12:21:18:21, no articles humped. Mouthparts unknown, space below for addition when described. Labrum [?broader than long, entire, rounded]. Mandibular incisor [?toothed, molar triturative, ratio of palp articles = 00:00:00, article 3 weakly falcate, setae = ABCDE]. Inner lobes of labium [?absent, weakly gaping]. Maxillae [?well setose medially, inner plate of maxilla 1 triangular, fully setose medially, outer plate with 7 spines, palps asymmetric]. Inner plate of maxilla 2 [?with oblique facial row of setae]. Outer plate of maxilliped [?medially spinose, article 3 of palp unlobed, dactyl shorter than 3, unguiform, with nail]. VOLUME 92, NUMBER 1 139 Coxae long, setae of medium size, coxa | curved forward, coxa 4 poorly (or strongly) lobed, coxa 5 shorter than 4. Gnathopods almost feeble, sub- chelate, almost of equal width, wrists of medium length, poorly lobed, hands subrectangular, palm oblique on gnathopod 1, transverse on gnathopod 2, short. Pereopods 3-7 not fossorial. Article 2 of pereopods S—6 scarcely expand- ed, tapering, moderately setose posteriorly or not, of pereopod 7 weakly expanded, unlobate or microlobate, moderately or strongly setose poste- riorly. Pleopods ordinary. Epimeron 3 lacking posteroventral fan of setae. Rami of uropods 1-2 extending subequally, peduncular setae absent, basofacial armaments [unknown], outer ramus of uropod 2 [possibly naked marginal- ly]. Uropod 3 scarcely extended, parviramous, outer ramus elongate, weak- ly setose and spinose, article 2 short. Telson of ordinary length, deeply cleft, lobes tapering, narrowed, weakly setospinose apically and with lateral setae. Gills [?2-6]. Oostegites [unknown]. Variants.—Inner ramus of uropod 3 slightly elongate (laeviuscula); oos- tegites narrow (brachyura); coxal gills 2-6, and ovate, some pediculate, (brachyura); coxa 4 slightly smaller than coxa 3 (in type-species, other species with normal coxa 4); gnathopod | distinctly larger than gnathopod 2 (brachyura); basis of pereopod 7 extremely expanded, almost lobate (brachyura); telson cleft only halfway (brachyura); cephalic sinus stronger (laeviuscula). Relationship.—Differing from Echinogammarus Stebbing in the curved coxa | and small, l-articulate accessory flagellum. Yogmelina ovata is es- pecially close to Echinogammarus warpachowskyi (Sars) because of the special shape on the basis of pereopod 7. But Y. ovata was said by Martynov to have a l-articulate accessory flagellum (warpachowskyi is said to have a 2-articulate accessory flagellum). Yogmelina brachyura is close to Baku paradoxus but differs in the 1- articulate accessory flagellum. See that genus for further diagnosis. Differing from Lanceogammarus andrussowi in the 1|-articulate accessory flagellum, the more equal gnathopods and the lack of strongly lobate bases on per- eopods 5-6. See Gmelina. Species.—brachyura (Derzhavin and Pjatakova, 1962), limana Karaman and Barnard, new species (=pusilla of Carausu, 1943); laeviuscula (Sars, 1896), ovata (Martynov, 1924), pusilla (Sars, 1896); Caspian and Black seas and their tributaries, 5. Yogmelina limana Karaman and Barnard, new species Gmelina pusilla.—Carausu, 1943:183-186, pls. 66, 67.—Carausu, Do- breanu, and Manolache, 1955:76, figs. 41-43 (not Sars, 1896). 140 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Name.—Referring to limans (estuaries) of Ponto Caspian deltas. Description of female. Body smooth, urosomites free, urosomites 1-2 with 2 setae and urosomite 3 with one seta on each dorsolateral surface. Rostrum short, lateral cephalic lobes protuberant, apically obtuse, antero- ventral sinus present. Eyes ovoid, as long as diameter of article 1 on an- tenna 1. Antenna | slightly shorter than antenna 2, ratio of peduncular articles = 28:20:14, relatively slender; accessory flagellum short, 1l-articulate, main flagellum with 8 articles. Antenna 2 ordinary; gland cone short, straight; article 3 short, article 4 as long as 5, flagellum 4-articulate, articles elongate. [Unknown = labrum, labium, maxilla 1, maxilla 2, maxilliped, all of man- dible except palp; pleopods, epimeron 1; oostegites, gills; males]. Article 1 of mandibular palp short, article 3 shorter than article 2, subfal- ciform, setae = ADE. Coxae of moderate length, 1—4 with long ventral setae; coxa | produced or curved forward, with concave anterior margin; coxa 4 scarcely lobed. Gnathopods small, subchelate, wrist gnathopod | shorter than hand, poorly lobed; gnathopod 2 scarcely larger than 1, wrist as long as hand, unlobed; hands of gnathopods 1-2 not alike, rectangular, palm well defined, oblique on gnathopod |, transverse on gnathopod 2. Pereopods 3-4 ordinary, posterior margin of pereopod 3 with sparse setae in bundles, article 5 very short. Pereopods 5-7; short basis of pereopods 5— 6 with parallel sides, with posterodistal lobe; basis of pereopod 7 expanded, with convex posterior margin, beveled distally, lobe absent; basis of per- eopods 6-7 setose mediofacially and of pereopods 5—7 on posterior margin. Epimera 2-3 angular, with long ventromarginal setae. Uropods 1 [?2] ordinary, rami extended subequally, spinose marginally, uropod 1 with groups of basofacial setae. Uropod 3 poorly extended, relatively short, par- Viramous, outer ramus longer than peduncle, article 2 short. Telson narrow, longer than wide, cleft three fourths, each lobe with 2 distal and 2 dorsofacial setae. Holotype.—Female (size unknown) figured by Carausu, 1943:183-186, plates 66, 67. Type-locality.—Lake Katlapug, on delta of Danube River, Black Sea. Relationship.—This species was identified as the Caspian species, Gmeli- na pusilla by Carausu but differs from that species in the lobed bases of pereopods 5-6, the unlobed basis of pereopod 7, the strongly setose epimera 2-3, and the presence of dorsal urosomal setae. Yogmelina limana differs from Y. ovata Martynov (1924, tributary of River Don) in the narrower and more poorly armed telson. The telson of Yogmelina ovata is very broad, tumid and heavily setose. The head, anten- nae, eyes, uropod 3 and pereopod 7 in part, correspond to those of Y. VOLUME 92, NUMBER 1 141 limana but otherwise Y. ovata is very poorly known and Y. limana, itself, is poorly known for many characters mentioned in the description above. Distribution.—Delta of Danube River. OTHER GAMMARIDAE Sandro, Karaman and Barnard, new genus Type-species.—Austroniphargus starmuhlneri Ruffo, 1960 (here select- ed). Name.—Named for the great Italian zoologist, Sandro Ruffo. Masculine. Body ordinary, smooth, urosomites coalesced. Rostrum and lateral ce- phalic lobes [unknown]. Eyes absent. Antennae elongate, antenna | longer than antenna 2, ratio of peduncular articles = [unknown], accessory flagellum l-articulate. Antenna 2 ordinary but article 1 grossly swollen. Labrum broader than long, entire, rounded. Mandibular incisor toothed, molar triturative, ratio of palp articles = 8:10:13, article 3 linear, setae = DE, but setae sparse and mostly near apex. Inner lobes of labium small, fleshy, well marked. Maxillae without medial setation (with medial hairs only), inner plate of maxilla 1 elongate-triangular, with 2 apical setae, outer plate with about 7 toothed spines (5 showing in illustration), palps asym- metric (left narrow and setose, right stout and spinose). Plates of maxilla 2 narrow, lacking medial and facial setae. Both plates of maxilliped of medium size, outer plate medially spinose, palp article 3 unlobate, dactyl as long as 3, unguiform, with nail. Coxae elongate, coxa 1 not dilated, coxa 4 lobate. Gnathopods of medium size, alike, wrist short, strongly lobed, hand trapezoidal, expanding apicad, palm weakly oblique, palm exceeding defining spines. Pereopods 3-4 ordinary. Pereopods 5-7 alike, of medium length, article 2 weakly expanded, weakly lobate, weakly setose posteriorly, posterior margins convex or weakly sinuous. Pleopods ordinary. Rami of uropods 1-2 extending subequally, marginally spinose, basofacial armaments [unknown]. Uropod 3 [?0f medium exten- sion], parviramous, peduncle with large apicolateral lobe, outer ramus elon- gate, moderately setose, article 2 short. Telson elongate, narrow, cleft three fourths, dorsally and distally spinose strongly. Coxal gills [?2-6], broadly ovate, pedicles [unknown]. Oostegites [un- known]. Relationship.—Allied to Austroniphargus Monod (1925:48) but differing in the presence of inner lobes on the labium, the shorter accessory flagellum, the longer coxae, the lobed coxa 4, the large lobe on the peduncle of uropod 3 and the normally developed pleopods. 142 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Species.—starmuhlneri (Ruffo, 1960); Madagascar, forest torrent but probably hypogean emergent, 1. Anopogammarus Derzhavin, revised Anopogammarus Derzhavin, 1945x: ?not seen.—Birstein and Levuschkin, 1970: 1478 (Anopogammarus birsteini Derzhavin, 1945x, ?monotypy). Body ordinary, smooth (type) or with 4 groups of strong spines on dorsal surface of metasomites (revazi); urosomites free. Rostrum short, lateral ce- phalic lobes acute, sinus present. Eyes absent. Antennae elongate, antenna | longer than 2, peduncular articles progres- sively shorter, primary flagellum elongate, accessory flagellum 4-articulate. Antenna 2 ordinary. Labrum broader than long, entire, rounded-truncate. Mandibular incisor toothed, molar triturative, ratio of palp articles = 6:19:15, article 3 weakly falcate, setae = BDE. Maxillae fully setose medially, inner plate of maxilla 1 triangular, fully setose medially, outer plate with 11 spines, palps asym- metrically armed, stout, slightly expanded distally, article 1 short. Inner plate of maxilla 2 with oblique facial row of setae. Outer plate of maxilliped medially spinose, articie 3 of palp unlobed, dactyl shorter than 3, unguiform, with nail. Coxae of moderate length, ventral margins lacking long setae; coxa 1 quadrate, coxa 4 lobate. Gnathopods large, almost identical but gnathopod 1 scarcely smaller than gnathopod 2, wrists short, lobed, hand Gammarus- like, elongate, palms almost identical, very oblique, long, with one spine near middle. Pereopods 3-4 ordinary. Pereopods 5-7 of medium length, almost of equal proportions, article 2 alike, scarcely expanded, tapering distally, posterior margin weakly sinuous, or weakly convex, with short setae, posteroventral corner weakly lobate (protuberant). Pleopods ordinary. Rami of uropods 1-2 extending equally, each ramus with marginal spine (possibly one absent in revasi), uropod 1 peduncle with basofacial spine [unknown in revazi]. Uropod 3 not extended, variramous, inner ramus reaching about halfway along article 1 of outer ramus, with few marginal setae or spines, outer ramus 2-articulate (type) or l-articulate (re- vazi). Telson of ordinary length, fully cleft, apices tapering, spinose, type with basolateral spines. Coxal gills 2-7, ovate to adze-shaped. Oostegites very broad. Relationship.—Differing from Echinogammarus Stebbing in the very weak lobation of pereopods 5-7. Differing from Zenkevitchia Birstein in the normal outer plate of maxilla 1 not grossly modified for filtration, in the large palp of maxilla 1, the normally short inner plate of the maxilliped; from Typhlogammarus Schaferna, Accubogammarus G. S. Karaman and VOLUME 92, NUMBER 1 143 Fontogammarus S. Karaman in the asymmetric palps of maxilla 1 bearing apical spines and from Metohia Absolon in the uncarinate body, variramous (not magniramous) uropod 3. I/vanella Vigna-Taglianti differs from Ano- pogammarus in the parviramous uropod 3. Anopogammarus revazi was originally described in Zenkevitchia but that genus is to be limited to species with the moplike maxilla 1 of filtering form in which dozens of spines occur on the outer plate and the palp is much reduced. Species.—birsteini Derzhavin, 1945, revazi (Birstein and Levuschkin, 1970); Transcaucasus, hypogean, 2. Tadzocrangonyx, Karaman and Barnard, new genus Type-species.—Crangonyx schizurus Birstein, 1948 (here selected). Name.—Crangonyx of Tadzhikistan. Neuter. Diagnosis.—Body ordinary, urosomites free, with several short, stiff dor- sal setae. Rostrum short, lateral cephalic lobes subrounded, anteroventral sinus weakly marked or absent. Eyes absent. Antenna 1 longer than antenna 2, ratio of peduncular articles = 21:10:9, accessory flagellum 2-articulate. Antenna 2 ordinary. Labrum entire, subrounded. Mandibular incisor toothed, molar tritura- tive, palp 3-articulate, article 1 short, 2 longer than 3, latter falciform (as in Gammarus). Labium ungaped, inner lobes weakly marked or absent. Max- illae 1-2 strongly setose medially, outer plate of maxilla 1 with 8-10 toothed spines, palps asymmetric. Inner plate of maxilla 2 with oblique facial row of setae. Plates of maxilliped of medium size, outer reaching one half to two thirds along palp article 2, palp articles 2-3 elongate, slender, article 3 un- lobed, dactyl unguiform, subapically setiferous. Coxae of medium size, coxa 1 unproduced, lobe of coxa 4 weak or absent. Gnathopods large, subchelate, gnathopod 2 the larger, wrist shorter than hand, weakly lobed, hand large, palms oblique, not spinose (spines only at defining or subdefining corners). Pereopods 3-4 ordinary. Pereopods 5-7 of medium size, 5 slightly the shortest, article 2 scarcely expanded basally, Gammarus-like, poorly or not lobed. Inner ramus of pleopods slightly to greatly shorter than outer ramus. Uropods 1-2 well developed, rami subequal to each other or inner ramus the shorter (schizurus); basofacial armaments [unknown]. Uropod 3 ex- ceeding apices of uropods 1-2, peduncle short, parviramous, inner ramus scale-like, outer ramus l-articulate, spinose. Telson of ordinary length, deeply incised, ungaped, apices strongly spinose. Coxal gills present, sternal gills absent. Oostegites [unknown]. Relationship.—Differing from Crangonyx Bate in the absence of bifurcate 144 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON spines and other spines on the palmar margins of the gnathopods (except at the defining corners), in the ungaped and deeply cleft telson, the higher number of spines on the outer plate of maxilla 1 (8-10 as opposed to 6-8), the slightly elongate articles 2-3 of the maxillipedal palp, and in the asym- metric palps of maxilla 1. The absence of sternal gills also occurs rarely in Crangonyx. Tadzocrangonyx bears a superficial resemblance to Protocrangonyx Nich- olls from Australia but differs in the absence of sternal gills, the incised telson, the higher number of spines on the outer plate of maxilla 1, the medially setose maxillae, and elongate uropod 3. Species.—setiferum (Birstein and Levuschkin, 1972) (Tjan-Shan, Middle Asia, USSR, from bank of Atbashi River); schizurum (Birstein 1948) (Ta- dzhikistan, Hissar, spring); probably both species are hypogean but emerge accidentally into epigean waters. Anisogammarus, Eogammarus, and Spinulogammarus Anisogammarus and Eogammarus are retained as valid genera but Spi- nulogammarus is synonymized with Eogammarus. Actually the differences between Anisogammarus and Eogammarus are not as strong as heretofore presented in the literature, the two genera clearly differing only in the pres- ence of a dorsal tooth or teeth on urosomites 1-2 in Anisogammarus and the absence of a tooth or teeth in Kogammarus. Evidence and discussion: The genera and species are as follows according to Tzvetkova (1975a); these references are omitted from our Literature Cited: 1. Anisogammarus Derzhavin, 1927, type-species: Anisogammarus dybov- skyi Derzhavin, 1927 (=Gammarus pugettensis Dana, 1853, =Gammarus pribilofensis Pearse, 1913); species: macginitei Shoemaker, 1955. 2. Eogammarus Birstein, 1933, type-species: Gammarus kygi Derzhavin, 1923 (selected by Tzvetkova, 1975); species: aestuariorum Tzvetkova, 1972; barbatus Tzvetkova, 1965; confervicolus Stimpson, 1856; hirsuti- manus Kurenkov and Mednikov, 1959; locustoides Brandt, 1851; mak- arovi Bulycheva, 1952; possjeticus Tzvetkova, 1967; ramellus Weckel, 1907; ryotoensis Ueno, 1940; schmidti Derzhavin, 1927; similimanus Bousfield, 1961; tiuschovi Derzhavin, 1927; turgimanus Shen, 1955. 3. Spinulogammarus Tzvetkova, 1972, type-species: Gammarus ochotensis Brandt, 1851; species: annandalei Tattersall, 1922; atchensis Brandt, 1851; jesoensis Schellenberg, 1937; oregonensis Shoemaker, 1944; spas- skii Bulycheva, 1952; subcarinatus Bate, 1862. Tzvetkova (1975a) considered these genera to be subgenera of Anisogam- MmMarus. VOLUME 92, NUMBER I 145 Anisogammarus contains species with a large erect tooth (knob or tuber- cle) on urosomite 2 or urosomites 1 and 2 whereas Eogammarus and Spi- nulogammarus each lack this ornamentation. The type-species of Aniso- gammarus has an elongate inner ramus on uropod 3 (magni- or variramous) and A. macginitei Shoemaker (1955) has a slightly elongate inner ramus. Tzvetkova illustrated specimens with an adequately variramous uropod 3 but Shoemaker’s original drawings show a much shorter inner ramus; we examined Shoemaker’s original material in the USNM and confirmed that the inner ramus is longer than the definition attributed to parviramous and that Tzvetkova is correct in attributing to the subgenus Anisogammarus third uropods with stronger inner ramus than the parviramous kind. How- ever, we must note that A. macginitei definitely represents an intergradation between parvi- and magniramous. We therefore find that the only incontest- able difference between Anisogammarus and Eogammarus remains the urosomal tooth (teeth) of Anisogammarus. Tzvetkova split the remaining species of the group, which theretofore had been assigned to Eogammarus, into 2 subgenera, Eogammarus and Spi- nulogammarus, based mainly on the presence of dorsal spination on the metasome in Spinulogammarus and its absence in Eogammarus. She also noted that most species of Eogammarus differed from most species of Spi- nulogammarus in the stronger setation of pereopods 5-7, uropod 3 and the telson, whereas in Spinulogammarus these appendages are furnished mainly with thick spines and few or no setae. The presence or absence of dorsal spination on the metasome has been abandoned as a good character in other genera such as Echinogammarus and though this is no justification for its abandonment in Eogammarus and Spinulogammarus, the species actually show transition and therefore the distinction is invalid. For example, several species of Spinulogammarus lack spines on metasomite | and several species of Eogammarus have weak to strong dorsal setae on metasomites 2-3, the difference between spines and setae being only degree of thickness. A transitional sequence follows: 1. S. annandalei bears setae or spines on the metasome or lacks any setae or spines on the metasome, all phases depending on age. 2. S. oregonensis, S. atchensis, S. ochotensis are fully spinose on meta- somites 1-3; a dorsal keel is present or absent. 3. S. subcarinatus is weakly spinose on metasomites 1-3 and also bears a dorsal keel. 4. S. jesoensis and S. spasskii lack spines on metasomite 1 but retain spines on metasomites 2-3. 5. E. ramellus bears setae on metasomites 2-3 and occasionally on meta- somite 1. 6. E. similimanus bears setae on metasomites 2-3. 146 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 7. E. confervicolus lacks metasomal setae and keel. 8. E. makarovi lacks metasomal setae but bears a Keel. Other species lack setae but the Keel is present or absent. None of the other potential generic characters distinctly separates the two genera as seen in the following sequences: Uropod 3. 1. Spines only: S. spasskii, S. ochotensis. 2. Spines and short setae: S. atchensis, E. makarovi, S. subcarinatus, S. annandalei. 3. Spines and longer or denser setae: S. oregonensis, E. confervicolus with setae increasingly longer or dense on E. kygi, E. tiuschovi, E. schmidti, E. barbatus, E. hirsutimanus, E. aestuariorum. Pereopod 7. 1. Setae absent: S. ochotensis, S. atchensis, S. subcarinatus, E. maka- rovi. 2. Setae sparsely present: E. aestuariorum. 3. Setae well developed: E. similimanus, E. ramellus, E. schmidti. Telson (in all cases spines are always present). 1. Setae absent: S. spasskii, S. subcarinatus, S. ochotensis, S. atchensis, E. ryotoensis, E. possjeticus. E. kygi, E. locustoides. 2. Setae weakly developed: S. oregonensis, S. annandalei, E. tiuschovi, E. barbatus, E. schmidti, etc. 3. Setae well developed: E. hirsutimanus, E. makarovi, E. ramellus, E. similimanus. Eogammarus Birstein, new status and new synonymy Eogammarus Birstein, 1933:149 (Gammarus kygi Derzhavin, 1923, selected by Tzvetkova, 1975a). (Spinulogammarus) Tzvetkova, in Golikov and Tzvetkova, 1972:2; Tzvet- kova 1972a:221; Tzvetkova 1972b:307 (Gammarus ochotensis Brandt, 1851, original designation). New synonym. Anisogammarus Derzhavin (newly restricted) Anisogammarus Derzhavin, 1927:8 (Gammarus pugettensis Dana, 1853 [=Anisogammarus dybovskyi Derzhavin, 1927, =Gammarus pribilofensis Pearse, 1913], monotypy). not Eogammarus Birstein, 1933:149 (=valid genus). not Spinulogammarus Tzvetkova, in Golikov and Tzvetkova, 1972:2 (=syn- onym of Eogammarus). Eriopisa Stebbing, revised Eriopis Bruzelius, 1859:64 (homonym, Insecta) (Eriopis elongata Bruzelius, 1859, monotypy). VOLUME 92, NUMBER 1 147 Eriopsis Wrzesniowsky, 1890:632 (spelling error). Eriopisa Stebbing, 1890:193 (replacement name); 1906:411 (same type- species). Body vermiform to subvermiform, urosomites free. Rostrum obsolescent, lateral cephalic lobes rounded, with deep thin sinus (notch). Eyes absent. Antennae elongate, antenna | much longer than antenna 2, ratio of pe- duncular articles = 24:26:8, accessory flagellum 2-articulate. Antenna 2 or- dinary but flagellum short, articles free. Labrum broader than long, entire or emarginate. Mandibular incisor toothed, molar triturative, ratio of palp articles = 4:11:16, article 3 linear, setae = ADE. Labium with small but fleshy inner lobes, gaping. Maxillae medially setose, inner plate of maxilla 1 ovatotriangular, fully setose me- dially, outer plate with 9 spines, palps almost symmetric. Inner plate of maxilla 2 with oblique facial row of setae. Outer plate of maxilliped medially setose, palp article 3 unlobed, dactyl about as long as 3, with nail. Coxae very short, often discontiguous, coxa 1 sharp anteriorly, coxa 4 unlobed. Gnathopods 1-2 dissimilar, subchelate, gnathopod 2 enlarged, wrist of gnathopod 1 of medium length, ovate, unlobed, hand trapezoidal, expanding distally, palm oblique, wrist of gnathopod 2 short, weakly lobed, hand enlarged, elongate, ovate, palm very oblique, long, poorly defined, often sculptured; female gnathopod 2 smaller than male, palm simple. Pereopods 3-4 ordinary. Pereopods 5-7 progressively longer but together not elongate, article 2 of pereopods 5-7 diverse, of 5 almost linear, of 6 slightly expanded, of 7 broadly expanded, 6-7 scarcely lobate. Pleopods ordinary. Rami of uropods 1-2 equally extended, with marginal spines, peduncle of uropod | with basofacial spine, of uropod 2 [?with ap- icomedial comb]. Uropod 3 greatly extended, parviramous, outer ramus huge, article 2 also huge and nearly as long as article 1. Telson of ordinary length to elongate, deeply cleft, narrow, lobes tapering to apical notch, poorly spinose or setose. Coxal gills 2-6, slender-ovate. Oostegites narrow. Notes.—We reexamined the type-species and found the inner lobes of the lower lip complete and fleshy (contrary to Sars, 1895, sketchy depiction). Relationship.—Differing from Melita in the elongate article 2 of the outer ramus on uropod 3, extremely thin body, and small coxae, with coxa 4 unlobed. All but type-species removed to genera following. Species.—elongata (Bruzelius, 1859) (Sars, 1895); bathyal north Atlantic and Pacific (some Pacific specimens may be distinct species or subspecies of this genus). Psammogammarus S. Karaman, revised Psammogammarus §S. Karaman, 1955:223 (Psammogammarus caecus S. Karaman, 1955, original designation). 148 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Body vermiform to subvermiform, urosomites free. Rostrum obsolescent or small, lateral cephalic lobes rounded, shallow, sinus obsolescent. Eyes absent. Antennae elongate, antenna 1 much longer than antenna 2, ratio of pe- duncular articles = 20:19:7, accessory flagellum 2-articulate. Antenna 2 or- dinary but flagellum short, articles free. Labrum broader than long, weakly emarginate. Mandibular incisor toothed, molar triturative, ratio of palp articles = 4:9:6, article 3 linear, setae = DE, very sparse. Labium with small but fleshy inner lobes partially fused together (like pleustids), gaping. Maxillae medially setose, inner plate of maxilla 1 ovate, not fully setose medially (3+ setae), outer plate with 9 spines, palps symmetric. Inner plate of maxilla 2 with oblique facial row of setae. Outer plate of maxilliped medially spinose (or only setose), palp ar- ticle 3 unlobed, dactyl as long as 3, unguiform, nail absent or obsolescent. Coxae very short, coxa | blunt anteriorly, coxa 4 unlobed. Gnathopods 1-2 subchelate, gnathopod 2 enlarged, wrist of gnathopod 1 of medium length, ovate, poorly lobed, hand weakly trapezoidal, weakly expanding apically, palm oblique, wrist of gnathopod 2 short, weakly lobed, hand en- larged, elongate, ovate, palm very oblique, poorly defined, often sculptured, female gnathopod 2 smaller than male, unsculptured, occasionally with weak hadziid setae. Pereopods 3-4 ordinary. Pereopods S-—7 progressively longer, last one elongate, article 2 of pereopods 5-7 alike, scarcely expanded, almost linear, scarcely to strongly lobate. Pleopods ordinary. Outer rami of uropods 1-2 slightly shortened, of uro- pod | lacking marginal spines, peduncle of uropod 1 with basofacial spine(s), of uropod 2 often with apicomedial comb. Uropod 3 highly extended, vari- ramous or parviramous, outer ramus hugely elongate, article 2 usually as long as article 1 but occasionally much shortened. Telson weakly elongate, deeply cleft, apices sharp, spinose, often with lateral spines, main dorsal setules strongly apicad. Coxal gills 3-6 on type-species, ovate to sausage-shaped. Oostegites [un- known on type-species]. Variants.—Species quite variable, for example: labrum rounded (philip- pensis), mandibular molar bulbous and poorly triturative (gracilis); article 3 of mandibular palp as long as article 2 (seurati); article 2 of antenna 1 shorter than article | (gracilis); lower lip normally melitid, with fully discrete fleshy inner lobes, no gape (/ongiramus); inner plate of maxilla 1 fully setose medially (ongiramus and seurati); nail of maxilliped discrete and outer plate of maxilliped only setose medially (philippensis); gnathopod 1 palm trans- verse (philippensis) or parachelate (seurati); article 2 of pereopods 3-4 ex- panded (philippensis); article 2 of pereopods 5-7 well lobate (philippensis); uropods 1-2 poorly spinose (gracilis); inner ramus of uropod 3 as long as VOLUME 92, NUMBER 1 149 article 1 of outer ramus (/Jongiramus) or half as long (caeca); article 2 on outer ramus of uropod 3 short (seurati); telson broadened (gracilis). Relationship.—Differing from Eriopisa in the lack of diversity on article 2 of pereopods 5-7 and the shorter article 3 of the mandibular palp, which in Eriopisa is much longer than article 2 and which in Psammogammarus is shorter than article 2. Species.—caecus S. Karaman, 1955 (=peresi Ledoyer, 1968), garthi (J. L. Barnard, 1952), gracilis (Ruffo and Schiecke, 1976), longiramus (Stock and Nijssen, 1965), philippensis (Chilton, 1921b), sewrati (Gauthier, 1936); cosmopolitan in low latitudes, anchialine or marine near brackish water, in wells or littoral, 6. Victoriopisa Karaman and Barnard, new genus Type-species.—Niphargus chilkensis Chilton, 1921a. Name.—Eriopisa of Victoria, Australia. Feminine. Body subvermiform, urosomites free, short. Rostrum very short, lateral cephalic lobes subrounded, with poorly marked anteroventral lobe, sinus obsolescent. Eyes weak or absent. Antennae elongate, antenna | longer than antenna 2, ratio of peduncular articles = 29:26:7 (or article 2 longer, epistomata); accessory flagellum 2-articulate. First article of flagellum on antenna 2 elongate and composed of several articles fused together, total free articles about 2-3. Labrum broader than long, entire, subrounded. Mandibular incisor toothed, molar triturative, ratio of palp articles = 4:11:16, article 3 linear, setae = DE. Inner lobes of labium partially fused together, obsolescent. Maxillae well setose medially, inner plate of maxilla 1 ovatotriangular, fully setose medially, outer plate with about 7 distal spines, palps [?symmetric][palp of epistomata shown to be l|-articulate]. Inner plate of maxilla 2 with several medial marginal setae only (none obliquely posi- tioned). Outer plate of maxilliped medially spinose, article 3 of palp unlobed, dactyl shorter than 3, unguiform, nail weak or absent. Coxae very short, discontiguous, coxa 1 not produced (or produced in epistomata), coxa 4 unlobed. Gnathopods 1-2 dissimilar, subchelate, gnath- opod 2 enlarged, wrist of gnathopod 1 elongate, ovate, unlobed, hand trap- ezoidal, expanding distally, palm almost transverse but convex; wrist of gnathopod 2 short, weakly lobed or not lobed, hand enlarged, elongate, ovate, palm very oblique, long, poorly defined, slightly sculptured, dactyl elongate. | Pereopods 3-4 ordinary. Pereopods 5-7 almost equal in length and short, article 2 of pereopods 5-7 diverse, of 5 unexpanded and almost linear, of 6 weakly expanded, of 7 broadly expanded and lobate; article 4 of pereopod 7 dilated (unusual character). 150 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Pleopods ordinary. Rami of uropods 1-2 subequally extended, with mar- ginal spines, peduncle of uropod | with basofacial spine, of uropod 2 with weak apicomedial comb. Uropod 3 greatly extended, parviramous, outer ramus huge, article 2 also huge and nearly as long as article 1. Telson of ordinary length, cleft to base, lobes tapering, weakly armed apically or apicolaterally. Coxal gills 2-6, some of them broadly pyriform. Oostegites [unknown]. Relationship.—Allied to Eriopisa Stebbing and Psammogammarus S. Karaman, but differing in the basal fusion of articles on the flagellum of antenna 2, the obsolescence of inner lobes on the labium, the absence of the oblique facial row of setae on maxilla 2, the dilated article 4 of pereopod 7, and the subequal pereopods 5 and 7. Species.—chilkensis (Chilton, 1921a) (Chilka Lake, India); australiensis (Chilton, 1923) (New South Wales, tidal lagoon) (Victoria); epistomata (Griffiths, 1974) (South Africa, open sea). Eriopisella sechellensis (Chevreux) Eriopisa sechellensis Chevreux, 1901:403, figs. 19-23.—Stebbing, 1906:732. Eriopisella sechellensis.—K. H. Barnard, 1935:284, fig. 4—Ruffo, 1959:6, fig. III, 1-2.—Nagata, 1965:304, fig. 33.—Sivaprakasam, 1969: fig. 3H—-K. Niphargus chilkensis.—Chilton, 1925:534, fig. 1 (not Chilton, 1921b). Article 2 of pereopod 7 varies from scarcely lobate in the typical material from the Seychelles Islands to moderately lobate in the Japanese material illustrated by Nagata. The material identified as a new subspecies by J. L. Barnard (1970), as E. sechellensis upolu has article 2 of pereopod 7 completely unlobed. We intend treating all subspecies as full species in our forthcoming monograph so that upolu is removed to specific level even though its distinctions are very small. Distribution.—Seychelles (type-locality); Japan to Red Sea, sublittoral. Giniphargus Karaman and Barnard, new genus Type-species.—Niphargus pulchellus Sayce, 1899. Name.—Niphargus of Gippsland. Masculine. Body subvermiform, urosomites free. Rostrum obsolescent, lateral ce- phalic lobes shallow, rounded, with sinus below and then bulbous corner extended. Eyes absent. Antennae slightly elongate, antenna | scarcely longer than antenna 2, ratio of peduncular articles = 30:24:9; accessory flagellum 4-articulate. Ar- ticle 4 of peduncle on antenna 2 slightly inflated, longer than article 5, fla- gellum ordinary, articles free. VOLUME 92, NUMBER I 151 Labrum broader than long, entire, rounded. Mandibular incisor toothed, molar triturative, ratio of palp articles = 10:18:15 (article 1 thus elongate), article 3 weakly falcate, setae = CDE. Inner lobes of labium discrete, small, fleshy. Maxillae moderately setose medially, inner plate of maxilla 1 ovate, with about 5 medial setae near apex, outer plate with about 8 serrate spines, palps [?symmetric]. Inner plate of maxilla 2 with oblique facial row of setae (as stated by description). Outer plate of maxilliped medially spinulate, ar- ticle 3 of palp unlobed, dactyl shorter than 3, unguiform, with nail. Coxae very short, barely touching each other, coxa 1 not produced, coxa 4 unlobed. Gnathopods 1-2 similar to each other, feeble, subchelate, mit- tenform, wrists unlobed, hands trapezoidal, small, expanding apically, palms transverse, short, wrist of gnathopod 1 of ordinary length, wrist of gnathopod 2 elongate, gnathopods otherwise identical in size. Pereopods 3-4 ordinary. Pereopods 5-7 progressively longer than each other but together not of elongate form, article 2 unexpanded, ovatolinear, unlobate, dactyls apparently with only one setule on inferior margin. Pleopods ordinary. Rami of uropods 1-2 extending subequally, marginally spinose, peduncle of uropod | apparently lacking basofacial spine. Uropod 3 greatly extended, of ‘‘parviramous’’ form because inner ramus absent, outer ramus huge, article 2 also huge and nearly as long as article 1. Telson short, broad, cleft halfway, apices broadly rounded, dorsally spinose. Coxal gills 2-6, ovoid, pedunculate, 2-articulate. Oostegites [unknown]. Relationship.—Differing from Indoniphargus Straskraba (1967) and Mi- croniphargus Schellenberg (1934) in the absence of a lobe on wrist of gnath- opod 1; from Eriopisella Chevreux in the absence of the inner ramus on uropod 3, the presence of an oblique setal row on maxilla 2, elongate article 2 on outer ramus of uropod 3, and, additionally, in the absence of a lobe on the wrist of gnathopod 2. Species.—pulchellus (Sayce, 1899); Australia, Victoria, Gippsland, pre- sumed hypogean, |. Pygocrangonyx Karaman and Barnard, new genus Type-species.—Metacrangonyx remyi Balazuc and Ruffo, 1953. Name.—Pygidized Crangonyx. Neuter. Body [?slender], urosomites free, naked. Rostrum obsolescent, lateral cephalic lobes subrounded, sinus [?present]. Eyes absent. Antennae elongate, antenna | longer than antenna 2, ratio of peduncular articles = 20:16:12, accessory flagellum [?2-articulate]. Antenna 2 ordinary. Labrum entire, broader than long, rounded apically. Mandibular incisor toothed, molar triturative, palp vestigial, l-articulate, setae = E. Labium without inner lobes. Maxillae 1-2 fully setose medially, inner plate of max- 152 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON illa 1 triangular, outer plate with 10 spines, palps asymmetrically armed. Inner plate of maxilla 2 with oblique facial row of setae. Both plates of maxilliped of medium size, outer plate [?medially spinose], palp article 3 unlobed, dactyl [?unguiform, ?with apical nail]. Coxae elongate, [?with short to medium setae], coxa | not expanded, coxa 4 unlobed, [?coxa 5 shorter than 4]. Gnathopods feeble, gnathopod 1 of Melita form, article 5 elongate, palm transverse; gnathopod 2 slightly larger, both articles 5 and 6 elongate and thin, palm very oblique, armed with bifid spines, posterior margin of hand with long curved hadziid-like setae. No sexual dimorphism. Pereopods 3-4 ordinary. Pereopod 7 slightly longer than pereopod 5, ar- ticle 2 of pereopods 5-7 lobate or not (7), weakly to strongly expanded, dactyls simple. Pleopods and uropods 1-2 ordinary, basofacial spine of uropod 1 [un- known]. Uropod 3 uniramous, very short, peduncle large, broader and lon- ger than ramus; inner ramus absent, outer ramus |-articulate, weakly longer than broad, stout, with 5 very long distal spines much longer than ramus itself. Telson very short, entire, broader than long, with 2 long distal spines longer than telson plus several setules. Coxal gills 2-6, narrow. Oostegites [unknown]. Relationship.—Alled to Metacrangonyx Chevreux (1909) but differing by the loss of the inner ramus on uropod 3, the much shortened outer ramus, the elongate distal spines on that ramus (1-2 shorter spines found in Me- tacrangonyx), the presence of long spines on the telson (setae in Metacran- gonyx) and by the absence of a lobe on article 2 of pereopod 7. These genera are not crangonyctids. Species.—remyi (Balazuc and Ruffo, 1953); Morocco, hypogean, 1. Dulichiella Stout, revived Dulichiella Stout, 1912:140 (Dulichiella spinosa Stout, 1912, monotypy). Body somewhat slender, urosomites free, metasomites and urosomites transversely crenulated or toothed dorsally. Rostrum short, lateral cephalic lobes subquadrate, sinus present. Eyes present. Antennae elongate, well setose, antenna | longer than antenna 2, ratio of peduncular articles = 16:18:4, ratio of flagella = 63:12, accessory flagellum multiarticulate (5-articulate in type). Ratio of peduncular articles 3,4,5 and flagellum on antenna 2 = 6:16:14:28. Labrum [‘‘slightly bilobed’’]. Mandibular incisor toothed, molar tritura- tive, ratio of palp articles = 3:8:10 (approximate), article 3 weakly clavate, setae of article 3 = ADE. Inner lobes of lower lip well developed, fleshy. Maxillary setae diverse; inner plate of maxilla 1 long, narrow, tapering, VOLUME 92, NUMBER 1 155 curved, with 1-2 strong apical plumose setae, medially margin hairy, outer plate with 9 spines, palp 2-articulate [symmetricity unknown]. Inner plate of maxilla 2 with oblique facial row of setae. Outer plate of maxilliped medially serrate and finely spinulose, article 3 of palp unlobed, dactyl un- guiform, nail weak. Coxae medium to short, moderately setose, coxa 1 scarcely expanded apically, coxa 4 poorly lobed and scarcely longer than coxa 5. Gnathopods subchelate, gnathopod 1 small, of Melita form, wrist elongate, unlobed, hand shorter than wrist, palm almost transverse; female gnathopod 2 slightly enlarged, wrist of same length as gnathopod 1 but hand much longer than wrist (thus wrist short), wrist scarcely lobate, palm weakly oblique, short, article 4 with tooth; one side (right or left) male gnathopod 2 like female gnathopod 2; other male gnathopod 2 (right or left) immensely enlarged and chelate, as in fiddler crabs, resembling crab claw, articles 3-5 tiny, hand immense, with giant chela, dactyl immense and thick, closing on transverse- ly extended palm. Pereopods 3-4 ordinary. Article 2 of pereopods 5-7 scarcely expanded, scarcely lobate ventrally. Pleopods ordinary. Rami of uropods 1-2 extending equally, marginally spinose, uropod | with basofacial spine on peduncle. Uropod 3 extended, parviramous, outer ramus elongate, article 2 short. Telson of ordinary length, cleft to base, but partially gaping, apically and laterally spinose. Coxal gills [?2-6, ovate]. Oostegites [?slender]. Relationship.—Like Melita but male gnathopod 2 of strong diversity from right to left sides, part of body with numerous transverse dorsal serrations, inner plate of maxilla 1 with sickle form and bearing only 1-2 fully apical setae. The species of this genus have been included with Melita since the late part of the 19th Century; Stout accidentally described this genus on the mistaken idea that uropod 3 lacked rami (because they had broken off) and thus compared the genus with Dulichia. Since Stout’s time the genus has always been synonymized with Melita but we believe the unusual male gnathopods analogous to those of a fiddler crab deserve generic recognition. In this way we also believe that the several species heretofore synonymized with Melita fresneli or M. appendiculata must be revived, reexamined and redescribed as there may be several valid species. For the moment, then, we revive the following species. Species.—appendiculata (Say, 1818); australis (Haswell, 1879) exilii (Fritz Muller, 1864); fresneli (Audouin, 1826) (?=cotesi Giles, 1890) (?=val- idus Dana, 1852) (=pilosus Dana, 1852) (=?setipes Dana, 1852), (=?aniso- chir Krgyer, 1845), (?=valida Dana, 1852); spinosa Stout, 1912; tropico- politan in shallow seas, ?5+. 154 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Quadrivisio Stebbing Quadrivisio bousfieldi, new species Quadrivisio bengalensis Bousfield, 1971:260-263, figs. 3, 4. Name.—Named for E. L. Bousfield, describer of the material. Accessory flagellum with 6-7 articles. Mandibular palp articles 2-3 ase- tose. Article 6 of male gnathopod | evenly rectangular. Epimera 1-3 with 1-2 posterior teeth each in addition to normal posterodistal tooth. Telson with 3 spines on each lobe but inner margins of lobes lacking armament. Holotype.—Female, ovigerous, 9 mm (figures 3—4 of Bousfield 1971). Type-locality.—Bismarck Archipelago, Manus Island, Liei River, outlet, 19 June 1962. Relationship.—This species differs from Q. bengalensis Stebbing, 1907 (and see Rabindranath, 1972; Nayar, 1959) by the absence of inner arma- ments on the telson, the absence of setae on distal palp articles of the mandible, and by the presence of 1-2 posterior teeth on epimera 1-3 in addition to the posterodistal tooth. Quadrivisio bousfieldi differs from Q. lutzi (Shoemaker, 1933) by the ab- sence of setae on the apex of the mandibular palp, the presence of super- numerary teeth on the evimera and in the shape of the lateral cephalic lobes. The new species differs from Q. aviceps K. H. Barnard (1940) in the supernumerary epimeral teeth and the presence of 3 spines (not 1) on each lobe of the telson. See the key to species below. Key to the Species of Quadrivisio° 1. Epimera 1-3 each with 1-2 posterior teeth besides normal poster- odistal tooth, palp articles 2-3 of mandible without setae ......... LeRe bas cintie sean hbiaddhs .otteuan bye Mitglied oye awe Qala aati ee ee QO. bousfieldi — Epimera 1-3 lacking supernumerary teeth, palp articles 2-3 of man- dible, with..1=2 setae each ccncsts secesdileQataot £ teres tae pa 2. Telson with spines or setae along inner margin of each lobe ...... sot Pores oleate cs, cite ad gh apitynuedl Bagh gah igtenaecieas ict NN Aen ee ee Q. bengalensis — Telson without spines or setae along inner margin of each lobe.... 3 3. Accessory flagellum with 11-12 articles; article 6 of gnathopod 1 in males dilated: distally.» sit Aco sers selene eee Q. aviceps — Accessory flagellum with 4-6 articles; article 6 of gnathopod 1 in males tapering distally... s,s. dheatss SOR eee Q. lutzi Nainaloa Karaman and Barnard, new genus Type-species.—Melita latimerus Bousfield, 1971. Name.—Contrived. Feminine. ” Quadrivisio chevreuxi Gordon and Monod (1968), is omitted because most of the key characters remain undescribed. VOLUME 92, NUMBER I 155 Body ordinary, smooth, urosomites free, urosomite 2 with 2 dorsal spines. Rostrum obsolescent, lateral cephalic lobes subrounded, prominent, antero- ventral sinus present. Eyes present. Antennae elongate, antenna | longer than antenna 2, peduncle almost slender, ratio of peduncular articles = 29:29:17; accessory flagellum 2-artic- ulate. Antenna 2 ordinary. Labrum entire, with weakly extended epistome in front. Mandibular in- cisor toothed, molar triturative, palp slender, 3-articulate, articles 2 and 3 subequally long, article 3 with 3 apical setae (E). Labium ordinary, with small fleshy inner lobes. Maxillae 1-2 not medially setose, inner plate of maxilla | with 3 apical setae, outer plate with 9 serrate spines, palp ordinary, 2-articulate, with distal setae and spines. Both plates of maxilla 2 narrow, inner lacking oblique facial row of setae. Both plates of maxilliped of me- dium size, outer plate setose along inner margin, palp article 2 elongate, article 3 short, slightly lobed, article 4 as long as article 3, unguiform, nail [unknown]. Coxae of medium size, coxa | not expanded, coxa 4 with shallow pos- terodistal lobe. Gnathopods strongly diverse, subchelate, sexually dimor- phic: article 5 of gnathopod 1 slightly longer than article 6, unlobed, palm transverse; gnathopod 2 enlarged, but weakly so in female, article 5 short to very short, with broad to narrow lobe, article 6 large, ovoid, palm strongly oblique, poorly defined. Pereopods 3-4 ordinary. Pereopods 5-7 not elongate, pereopod 5 weakly shorter than pereopod 7, article 2 of all expanded, ovoid, lobate. Pleopods weak but rami multiarticulate. Uropod | slightly reduced, [?ap- parently without basofacial spine], outer ramus slender, naked dorsally, inner ramus slightly elongate, dorsaily spinose. Uropod 2 ordinary, reaching as far as uropod 1, rami normally spinose. Uropod 3 strongly exceeding uropod 1, parviramous, peduncle shorter than outer ramus, inner ramus scale-like, outer ramus rectangular, with medium article 2. Telson short, cleft to base, each lobe quadrate, weakly excavate apically, weakly spinose. Gills simple, 2-6, ovate to linear, not strongly pediculate. Oostegites nar- row. Relationship.—Allied to Rotomelita J. L. Barnard (1977), but differing from it in the partly reduced uropod 1, subequally long pereopods 5-7, simple gills, sexual dimorphism of gnathopod 2, lobed article 2 of pereopods 5-7, presence of eyes and prominent lateral cephalic lobes. Species.—latimerus eae acins 1971); Bismarck Archipelago, brackish lakes and lagoons. Gammarella Bate, new synonymy Pherusa Leach, 1814:432 (homonym, Polychaeta) (Pherusa fucicola Leach, 1814, monotypy). 156 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Gammarella Bate, 1857:143.—Stebbing, 1906:449 (Gammarella orchesti- formis 1857, monotypy, =Pherusa fucicola Leach). Pherusana J. L. Barnard, 1964:62 (new name for Pherusa, same type- species). Nuuanu J. L. Barnard, 1970:166 (Nuuanu amikai J. L. Barnard, 1970, orig- inal designation). New synonym. Cottesloe J. L. Barnard, 1974:27 (Cottesloe berringar J. L. Barnard, 1974, original designation). New synonym. Body laterally compressed, carinate or smooth, urosomites free, carinate or smooth, weakly spinose, urosomite 2 often with pair of dorsal spines. Rostrum short, lateral cephalic lobes acute at upper corner but actually forming large quadrate extension with narrow incision. Eyes present but often poorly developed. Antennae elongate to medium, joints often geniculate, peduncular article 1 slender to stout, article 2 longer or shorter than article 1, article 3 shorter than 2; typical ratio of articles = 26:30:10, accessory flagellum 3+-articu- late. Antenna 2 thin, shorter than antenna 1. Labrum broader than long, subrounded, entire. Mandibular incisor toothed, molar triturative, palp article 1 weakly to strongly elongate, article 2 slender, elongate, article 3 falcate, shorter than or equal to article 2, typical ratio = 5:14:11, setae = DE. Inner lobes of labium absent or faintly marked. Maxillae strongly setose medially, inner plate of maxilla 1 leaf-like or subtriangular, heavily setose medially, outer plate with 9 spines, palps asymmetric, 2-articulate. Inner plate of maxilla 2 with oblique facial row of setae. Maxillipedal plates large, outer medially spinose, palp strong, article 3 unlobed, dactyl shorter than 3, unguiform, with weak nail. Anterior coxae of medium extension, or long, coxa | scarcely expanded apically, coxa 4 lobed (coxa 3 rarely shortened), coxa 5 shorter than 4. Gnathopods subchelate, of female almost feeble, dissimilar, almost of equal size, wrists elongate, unlobed, hands narrow, rectangular or subrectangular, palm of gnathopod | usually transverse, of gnathopod 2 oblique and often obsolescent; male gnathopod 1 like female, male gnathopod 2 greatly en- larged, wrist very short, cryptic (type) or strongly lobed, hand elongate, ovate, palm oblique and long or merging with posterior margin, dactyl elon- gate, curved, or short and stout. Pereopods 3-4 ordinary. Article 2 of pereopods 5—7 usually strongly ser- rate posteriorly (but not in type), otherwise dissimilar in shape, of pereopods 5—6 weakly expanded, tapering distally, moderately lobate posteroventrally, of pereopod 7 broadly expanded, shield-like, these pereopods usually short, pereopod 6 often slightly the longest. Pleopods ordinary. Rami of uropods 1-2 extending equally, spinose mar- ginally, uropod 1 with basofacial spine. Uropod 3 not extended, parvira- VOLUME 92, NUMBER 1 ee / mous, inner ramus occasionally however with medial spine(s), outer ramus short (thus making inner ramus appear much larger than it is relative to genera such as Melita), spinose, article 2 small (possibly absent in male of type-species). Telson short, deeply cleft, lobes tapering sparsely, spinose apically. Coxal gills 2-6, ovate, not pediculate. Oostegites slender. Variants.—Articles of peduncle on antenna | often fixed in geniculate fashion in preserved material (especially species of Nuuanu); article 2 of antenna | varying between 0.55 and 1.25 times length of article 1; article 2 of pereopods 5-7 scarcely serrate (fucicola), or strongly serrate (all other species); urosomite | with dorsal tooth (fucicola, berringar) or not (most other species); body cuticle with straw-setules (fucicola, species of Nuuanu) or villose (berringar, merringannee). Remarks.—When Nuuanu was originally described, its affinity with Gam- marella was overlooked because of the much more strongly developed ex- pansions and serrations on pereopods 5-7 and the strange head shape not previously noted for Gammarella except by Sowinsky (1898) and overlooked by Barnard. Then Cottesloe was established with affinities to Nuuwanu but strong differences in antenna | article ratios, cuticle texture but especially robust body size and opaque cuticle (latter as seen in retrospect). Finally, Tabatzius McKinney and Barnard (1977) was described on the basis of parasitic maxillae and a specimen of Gammarella reappraised. We have now examined more specimens of G. fucicola and have concluded, despite a wide variety of urosomal teeth, cuticles, antennae, eyes, robust bodies, pereopodal shapes and serrations, that no discontiguity exists among Gam- marella, Nuuanu and Cottesloe. One species, Nuuanu mokari J. L. Barnard (1974) has a much shortened coxa 3 and could be elevated to generic status but we reserve this action until more exploration in the IndoPacific dis- proves any intergradation for that character. Meanwhile Tabatzius is re- tained but without strong conviction as it may also be found to have inter- gradational relatives yet undiscovered. Species.—amikai (J. L. Barnard, 1970), berringar (J. L. Barnard, 1974), fucicola (Leach, 1814) (Sowinsky, 1898; Chevreux and Fage, 1925, but not well depicted), merringannee (J. L. Barnard, 1974), mokari (J. L. Barnard, 1974), numbadi (J. L. Barnard, 1974); eastern Atlantic, Mediterranean, Indo-Pacific especially Australia and Hawaii (as yet explored), 6. Tabatzius McKinney and Barnard, emended Tabatzius McKinney and Barnard, 1977:163 (Tabatzius copillius McKinney and Barnard, 1977, original designation, =Nuuanu muelleri Ortiz, 1976). Tabatzius muelleri (Ortiz), new combination, new synonymy 158 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Nuuanu muelleri Ortiz, 1976:13, figs. 1-3. Tabatzius copillius McKinney and Barnard, 1977:164, figs. 1-3. New syn- onym. : Remarks.—The name proposed by Ortiz takes priority over T. copillius. In McKinney and Barnard (1977, fig. 3) the middle gnathopod labeled ‘*G2”’ should be labeled “‘G2c.”’ Afridiella Karaman and Barnard, new genus Type-species.—Bogidiella somala Ruffo, 1970. Name.—Contrived, from Africa. Feminine. Body [?subvermiform], urosomites [?free]. Rostrum obsolescent, lateral cephalic lobes [?subrounded, ?sinus present]. Eyes absent. Antennae elongate, antenna | longer than 2, ratio of peduncular articles = 23:22:12, accessory flagellum 2-articulate, primary flagellum slightly shorter than peduncle. Antenna 2 ordinary. Labrum [?entire, rounded]. Mandibular incisor weakly toothed, connect- ed to large excavate callus, molar small, conical, not triturative, with strong comb and long seta, ratio of palp articles = 7:12:6, article 3 linear, setae = E. Labium [?almost gaping, inner lobes fused together]. Maxillae not me- dially setose, inner plate of maxilla 1 onion-shaped, with 2 apical setae, outer plate with 7 spines, palp apically slender, sparsely setose. Inner plate of maxilla 2 lacking facial and medial setae. Plates of maxilliped small, outer medially setose very sparsely, palp strong, article 3 unlobed, dactyl as long as 3, unguiform, with nail. Coxae short, but longer than broad, coxa 4 unlobed, coxa 5 [?as long as 4 and rather large]. Gnathopods large, gnathopod 1 scarcely larger than 2, wrists short to medium respectively, strongly to moderately lobed respec- tively, hands elongate, slightly larger on gnathopod 1, palms very oblique, lined with short setae, poorly defined, somewhat longer on gnathopod 2, dactyls elongate. Pereopods 3-4 ordinary, article 2 not dilated. Article 2 of pereopod 5 almost rectilinear, weakly expanded on pereopods 6—7, unlobate, dactyls short; article 6 of pereopods 5-7 anteriorly setose. Pleopods with vestigial inner ramus, outer ramus short, 3-articulate. Rami of uropods 1-2 extending equally, without marginal spines, peduncle of uropod | with basofacial or ventrolateral spines. Uropod 3 extended, mag- niramous, rami elongate, l-articulate, almost aequiramous. Telson short, broad, scarcely emarginate, with distal and apicolateral spines on each side. Coxal gills 4-6, ovate. Oostegites small, geniculate, dense setae confined apically. Relationship.—Differing from Bogidiella Hertzog (1933) in the excavate mandibular callosity, the strongly spinose molar, elongate palp article 1 of VOLUME 92, NUMBER 1 159 the mandible, and slightly enlarged coxae. See following key for generic position. Species.—somala (Ruffo, 1970); Somalia, hypogean, 1. Key to the Genera of *‘Bogidiellids”’ fPMROMOPOC St UMIGATMOUS). soe ce be eecee ye autey ate. cs ays sy bieiae 3 * Pseudingolfiella Sn eC N Ge OME ANT OUS ie ecni Paha seudthe Chia uch 2) cs toead aac So sleslsuchyoy Slee ity whales due. 2 PWKOPOG.5 PALVIFAMOUS ser4)o vations dasnieeanay. wae de evar Paracrangonyx See POpPOdsS Aad OMINATITOUS MMe ene ek EL PR AE EL LPS. 3 SECO POU HeUNIPAMOUS Ss Che ene SRE EEL UCN Po Bollegidia SRLS RON OG Me MG ATMOUS i cp ee ce ag ene oe oa ges ops con 3 ko cain a mo oye § 4 AveOutenr rami, Ol PIEOPOS, P-artiCulAate i165 .cues ea: op Sparonerdhons crane Kergueleniola Ouch! Ofpleopods multianticulaters i... 6) vanced ains sO. os 5 5. Coxae shorter than broad, often discontiguous or barely touching . ee eT nee PAR eC LM On eee Tes 5 4 ss 6 — Coxae longer than broad, several coxae strongly overlapping ..... so 3b a ea iy ae ee ee Onn fare Spelaeogammarus 6. Mandibular incisor ordinary, molar weakly to strongly triturative, NOUSPINGSe:onmpoorly Spinose fate FA Bogidiella — Mandibular incisor with large excavate callosity, molar not triturative, SLPOMGLNG GOWN CSS ae sale eye eee eI a CAN Afridiella Literature Cited Audouin, V. 1826. Explication sommaire des plances de Crustacés de l’Egypte et de la Syrie, publiées par Jules César Savigny, membre de |’Institut; offrant un exposé des caractéres naturels des genres, avec la distinction des espéces.—Description de l’Egypte, Histoire Naturelle, 1:77-98. Balazuc, J., and S. Ruffo. 1953. Due nuove species del genre Metacrangonyx Chevreux (Amphipoda-Gammaridae) delle acque interne del Nord Africa Francese.—Memorie del Museo Civico di Storia Naturale Verona 4:25-33, figs. 14. Barnard, J. L. 1952. A new species of amphipod from Lower California (Genus Eriopisa).— Pacific Science 6:295-—299, figs. 1-2. . 1954. Marine Amphipoda of Oregon.—Oregon State Monographs, Studies in Zoology 8:1-103, 1 unnumbered fig, pls. 1-33. . 1964. Revision of some families, genera and species of gammaridean Amphipoda.— Crustaceana 7:49-74. . 1969a. The families and genera of marine gammaridean Amphipoda.—United States National Museum Bulletin 271:1—535, figs. 1-173. . 1969b. Gammaridean Amphipoda of the rocky intertidal of California: Monterey Bay to La Jolla—United States National Museum Bulletin 258: 1-230, figs. 1-65. . 1970. Sublittoral Gammaridea (Amphipoda) of the Hawaiian Islands.—Smithsonian Contributions to Zoology 34: 1-286, figs. 1-180. . 1971. Gammaridean Amphipoda from a deep-sea transect off Oregon.—Smithsonian Contributions to Zoology 61:1-86, figs. 148. . 1972. The marine fauna of New Zealand: Algae-living littoral Gammaridea (Crustacea Amphipoda).—New Zealand Oceanographic Institute Memoir 62:7-216, figs. 1-109. 160 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON . 1974. Gammaridean Amphipoda of Australia, part I1.—Smithsonian Contributions to Zoology 139:1-148, figs. 1-83. . 1977. The cavernicolous fauna of Hawaiian lava tubes 9. Amphipoda (Crustacea) from brackish lava ponds on Hawaii and Maui.—Pacific Insects 17:267-299, figs. 1-16. 1978. Redescription of the amphipod, Calliopiella michaelseni Schellenberg from South Africa, with comparison to a new genus from the Pacific Ocean.—Annals of the South African Museum 75:33-40, figs. 1-3. Barnard, K. H. 1930. Amphipoda.—British Antarctic (“Terra Nova’’) Expedition, 1910, Nat- ural History Reports, Zoology 8:307-454, figs. 1-63. . 1931. Diagnosis of new genera and species of amphipod Crustacea collected during the ‘‘Discovery”’ Investigations, 1925—1927.—Annals and Magazine of Natural History, series 10, 7:425-430. . 1932. Amphipoda. Discovery Reports 5: 1-326, figs. 1-174, pl. 1. . 1935. Report on some Amphipoda, Isopoda and Tanaidacea in the collections of the Indian Museum.—Records of the Indian Museum 37:279-319, figs. 1-21. . 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 the hitherto recorded marine and fresh-water species.—Annals of the South African Museum 32:381-543, figs. 1-35. 1955. Additions to the fauna-list of South African Crustacea and Pycnogonida.— Annals of the South African Museum 43:1-107, figs. 1-53. Bate, C. S. 1857. A synopsis of the British edriophthalmous Crustacea.—Annals and Magazine of Natural History, series 2, 19:135-152, figs. 1-2. . 1858. 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De skandinaviske og arktiske Amphipoder.—A. W. Brggger, Christiania:i— IVa t= 2eeplswal=32- Bousfield, E. 1971. Amphipoda of the Bismarck Archipelago and adjacent Indo-Pacific islands (Crustacea).—Steenstrupia 1:255-293, figs. 1-20. Bruzelius, R. 1859. Bidrag till kannedomen om Skandinaviens Amphipoda Gammaridea.—K. Svenska Vetenskapsakademien Handlingar, new series, 3:1-104, pls. 1-4. Bryazgin, V. F. 1974. A contribution to the fauna of Gammaridea (Amphipoda) in the Barentz Sea.—Akademiia Nauk SSSR, Zoologicheskii Zhurnal, 53:1417-1420, figs. 1-4. Bulycheva, A. I. 1952. Novye vidy bokoplavov (Amphipoda, Gammaridea) iz Japonskogo Morja.—Akademiia Nauk SSSR, Trudy Zoologicheskogo Instituta, 12:195-250, figs. 1-39. Carausu, S. 1943. Amphipodes de Roumanie I. Gammaridés de type Caspien.—Institutul de Cercetari Piscicole al Romaniei, Monographia 1:1-293, figs. 1-20, pls. 1-85. VOLUME 92, NUMBER 1 161 , E. Dobreanu, and C. Manolache. 1955. 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Conspectus crustaceorum quae in orbis terrarum circumnavigatione, Carolo Wilkes e classe reipublicae faederatae duce, lexit et descripsit. Pars III. (Amphipoda.No.I).—American Academy of Arts and Science, Proceedings 2:201-—220. . 1853. Crustacea. Part II—U.S. Exploring Expedition 14:689-1618. Atlas of plates 1- 96 [published in 1855]. Dediu, I. I. 1967. Amfipody i mizidy basseinov rek Dnestra i Pruta. Sistematika, ekologiia, zoogeograficheskii analiz i khoziaistvennoe znachenie.—Akademiia Nauk Moldavskoi SSR, Institut Zoologii. Izdatel’stvo ‘‘Nauka’’ Moskva 1967:1-171, figs. 1-19. Della Valle, A. 1893. Gammarini del Golfo di Napoli—Fauna und Flora des Golfes von Neapel und der angrenzenden Meeres-Abschnitte, Monographie 20:i—xi, 1-948, atlas of plates 1-61. Derzhavin, A. N., and G. M. Pjatakova. 1962. Novye vidy Kaspiiskikh amfipod.—Akademiia Nauk Azerbaidzhanskoi SSR, Doklady 18:53-57, pls. 1-2. . 1967. Novye vidy gammarid Kaspiiskogo Morja, pages 79-84, figures 1-2, in: M. A. Musaev et alia, editors ‘‘Biologicheskaja Produktivnost’’ Kurinsko-Kaspiiskogo Ry- boloviogo Raiona, Akademija Nauk Azerbaidzhanskoi SSR, Institut Zoologii. Dunbar, M. J. 1954. The amphipod Crustacea of Ungava Bay, Canadian Eastern Arctic.— Journal of the Fisheries Research Board of Canada 11:709-798, figs. 1-42. Fabricius, J. Ch. 1779. Reise nach Norwegen mit Bemerkungen aus der Naturhistorie und Oekonomie.—Hamburg, Carl Ernst Bohn. Gauthier, H. 1936. Eriopisa seurati, nouvel amphipode du Sud-Tunisien.—Bulletin de la So- ciété d’Histoire Naturelle de |’ Afrique du Nord 27:133-143, figs. 1-3. 162 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Giles, G. 1890. Descriptions of seven additional new Indian amphipods. Natural History notes from H. M. Indian Marine Survey Steamer ‘‘Investigator,’’» Commander Alfred Car- penter, R.N., D.S.O., commanding. -No.15.—Journal of the Asiatic Society of Bengal 59:63-74, pl. 2. Gordon, I., and Th. Monod. 1968. Sur quelques Crustacés des eaux douces de Zanzibar.— Bulletin de Il’ Institut Fondamental d’ Afrique Noire, série A, Sciences Naturelles 30:497-— 517, figs. 1-31. Griffiths, C. L. 1974. The Amphipoda of Southern Africa Part 2. The Gammaridea and Ca- prellidea of South West Africa south of 20°S.—Annals of the South African Museum 62: 169-208, figs. 1-7. . 1975. The Amphipoda of Southern Africa Part 5. The Gammaridea and Caprellidea of the Cape Province west of Cape Agulhas.—Annals of the South African Museum 67:91-181, figs. 1-21. Gurjanova, E. F. 1938. Amphipoda, Gammaroidea of Siaukhu Bay and Sudzukhe Bay (Japan Sea).—Reports of the Japan Sea Hydrobiological Expedition of the Zoological Institute of the Academy of Sciences, USSR, in 1934, 1:241-404, figs. 1-59. . 1972. Novye vidy bokoplavov (Amphipoda, Gammaridea) iz severozapadnoi chasti Tikhogo Okeana.—Akademiia Nauk SSSR, Trudy Zoologicheskogo Instituta 52:129- 200, figs. 1-43. Haswell, W. A. 1879. On Australian Amphipoda.—Proceedings of the Linnean Society of New South Wales 4:245-279, pls. 7-12. Hertzog, L. 1933. Bogidiella albertimagni sp.nov., ein neuer Grundwasseramphipode aus der Rheinebene bei Strassburg.—Zoologischer Anzeiger 102:225—227, fig. 1. Hurley, D. 1954. Studies on the New Zealand amphipodan fauna. No. 9. The families Acan- thonotozomatidae, Pardaliscidae and Liljeborgiidae.—Transactions of the Royal Society of New Zealand 82:763-802, figs. 1-14. Just, J. 1977. Amphyllodomus incurvaria gen.et sp.n. (Crustacea, Amphipoda), a remarkable leaf-cutting amphithoid from the marine shallows of Barbados.—Zoologica Scripta 6:299-332, figs. 1-3. Karaman, S. 1931. II. Beitrag zur Kenntnis der Amphipoden Jugoslaviens, sowie einiger Arten aus Griechenland.—Prirodoslovne Razprave, Ljubljana 1:31-66, figs. 1-11. . 1955. Uber einige Amphipoden des Grundwassers der Jugoslavischen Meereskiiste — Acta Musei Macedonici Scientiarum Naturalium, Skopje 2:223-242, figs. 1-51. Krgyer, H. 1845. Karcinologiske Bidrag.—Naturhistorisk Tidsskrift 1:283-345, pls. 1-3; 403, 453-638, pls. 6-7. ——. 1846. Karcinologiske Bidrag (Fortsaettelse) —Naturhistorisk Tidsskrift 2:1-211, pls. 1-2. Kunkel, B. 1910. The Amphipoda of Bermuda.—Transactions of the Connecticut Academy of Arts and Sciences 16:1-116, figs. 1-43. Leach, W. E. 1814. Crustaceology. Appendix.—Edinburgh Encyclopaedia 7:429-434. Ledoyer, M. 1968. Ecologie de la faune vagile des biotopes Méditerranéens accessibles in scaphandre autonome (région de Marseille principalement). IV.-Synthése de l'étude écologique.—Recueil des Travaux de la Station Marine d’Endoume, Bulletin 44, 60:125— 295, pls. 1-24. ———. 1972. Amphipodes gammariens vivant dans les alvéoles des constructions organogenes récifales intertidales de la région de Tuléar (Madagascar). Etude systématique et éco- logique.—Tethys Supplement 3:165—285, figs. 1-2, pls. 1-80. ———. 1973. Etude des amphipodes gammariens des biotopes de substrats sableux et sablo- vaseux de la région de Tuléar et de Nosy-Bé (Madagascar).—Tethys Supplement 5:51- 94, pls. 1-30. Martynov, A. 1919. Sur les Crustacés superieurs des environs du Rostov sur Don.—Arbeiten der Naturf.Ges.d.Donnschen Univ. Rostov 1:39-53 [not seen]. VOLUME 92, NUMBER 1 163 1924. Etudes sur les Crustacées de mer du bassin du bas Don et leur distribution éthologique.—Akademii Nauk SSSR, Leningrad, Ezhegodnik Zoologicheskogo Museja 25 A =115; spl. McCain, J. 1971. A new deep-sea species of Epimeria (Amphipoda, Paramphithoidae) from Oregon.—Crustaceana 20: 159-166, figs. 1-3. McKinney, L., and J. L. Barnard. 1977. A new marine genus and species of the Nuuanu- group (Crustacea, Amphipoda) from the Yucatan Peninsula.—Proceedings of the Bio- logical Society of Washington 90:161-171, figs. 1-3. Monod, Th. 1925. Niphargopsis bryophilus et var. petiti, gen., sp. et var. nov., amphipode nouveau des eaux douces de Madagascar.—Bulletin de la Société Zoologique de France 50:40-48, figs. 1-3. Mordukhai-Boltovskoi, F. D., and S. M. Ljakhov. 1972. Novyi vid amfipod roda Stenogam- marus (Gammaridae) v basseine Volga.—Zoologicheskii Zhurnal 51:21—27, figs. 1-2. Muller, Fritz. 1864. Fur Darwin.—Leipzig, Wilhelm Engelmann, 91 pages, 65 figs. Nagata, K. 1965. Studies on marine gammaridean Amphipoda of the Seto Inland Sea. HI.— Publications of the Seto Marine Biological Laboratory 13:291-326, figs. 27-44. Nayar, K. N. 1959. The Amphipoda of the Madras Coast.—Bulletin of the Madras Govern- ment Museum, new series, Natural History Section 6(3):1—-59, pls. 1-16. Ortiz, M. 1976. Un nuevo genero y una nueva especie de anfipodo de aguas Cubanas (Am- phipoda, Gammaridea, Ampithoidae).—Ciencias, Universidad de la Habana, Investi- gaciones Marinas 27:1—26, 2 figs. [combination of 3 papers]. Rabindranath, P. 1972. Studies on Gammaridean Amphipoda (Crustacea) from India.—Bul- letin Zoologisch Museum, Universiteit van Amsterdam 2:155-172, figs. 1-7. Rathke, H. 1843. Beitrage zur Fauna Norwegens.—Verhandlungen Kaiserlichen Leopoldi- nisch-Carolinischen Akademie Naturforscher, Breslau, 20:1—264, pls. 1-12. Ross, J. C. 1835. Zoology in: Appendix to the narrative of a second voyage in search of a North-West Passage, and of a residence in the arctic regions during the years 1829, 1830, 1831, 1832, 1833. By Sir John Ross, C. B., K. S. A., K. C. S., &c. &c. Captain in the Royal Navy. Including the reports of Commander now Captain, James Clark Ross, R.N., F.R.S., F.L.S., &c. and the discovery of the northern magnetic pole. London, A.W. Webster:vii-c, pls. A-C. Ruffo, S. 1959. Contributo alla conoscenza degli anfipodi del Mar Rosso (1).—Sea Fisheries Research Station, Haifa, Bulletin 20: 1-36, figs. 1-6. 1960. Studi sui crostacei anfipodi LVI Anfipodi raccolti dalla spedizione austriaca 1958 nel Madagascar con descrizione di una nuova specie di Austroniphargus Monod.— Mémoires de |’Institut Scientifique de Madagascar, série A, 14:63-69, figs. 1-2. . 1970. Studi sui crostacei anfipodi LXIV. Bogidiella somala n.sp. delle acque sotter- ranee della Somalia (Crustacea Amphipoda).—Monitore Zoologico Italiano, NS supple- mento III:159-171, figs. 1-5. , and U. Schiecke. 1976. Descrizione di Eriopisa gracilis n.sp. (Amphipoda, Gammar- idae) delle Coste di Malta e redescrizione di E. coeca (S. Karaman, 1955) (=E. peresi M. Ledoyer, 1968).—Bollettino del Museo Civico di Storia Naturale Verona 2:415—438, figs. 1-8. Sars, G. O. 1876. Prodromus descriptionis Crustaceorum et Pycnogonidarum, qvae in expe- ditione Norvegica anno 1876, observavit.—Archiv for Mathematik og Naturvidenskab DDD le . 1894. Crustacea Caspia. Contributions to the knowledge of the carcinological fauna of the Caspian Sea. Part HI. Amphipoda. Ist Article, Gammaridae (part).—Bulletin de l’ Académie Impériale des Sciences de St.-Pétersbourg, série 5, 1:179—223, pls. 1-8. . 1894a. Crustacea Caspia. Contributions to the knowledge of the carcinological fauna of the Caspian Sea. Part III. Amphipoda. Second Article: Gammaridae (continued).— 164 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Bulletin de 1’ Académie Impériale des Sciences de St.-Petersbourg, série 5, 1:343-378, pls. 9-16. —. 1895. Amphipoda. An account of the Crustacea of Norway with short descriptions and figures of all the species, 1:i—viii, 1-711, pls. 1-240, 8 supplementary pls. . 1895a. Crustacea Caspia. Contributions to the knowledge of the carcinological fauna of the Caspian Sea. Part III. Amphipoda. Third Article. Gammaridae (concluded). Cor- ophiidae.—Bulletin de |’ Académie Imperiale des Sciences de St.-Petersbourg, série 5, 3:275-314, pls. 17-24. . 1896. Crustacea Caspia. Contributions to the knowledge of the carcinological fauna of the Caspian Sea. Amphipoda, Supplement.—Bulletin de | Academie Impériale des Sciences de St.-Pétersbourg, série 5, 4:421-489, pls. 1-12. Say, Th. 1818. An account of the Crustacea of the United States.—Journal of the Academy of Natural Sciences of Philadelphia 1:374-401. Sayce, O. A. 1899. Niphargus pulchellus, a new Victorian blind amphipod.—Proceedings of the Royal Society of Victoria, new series 12:152-159, pls. 15, 16. Schellenberg, A. 1926. Die Gammariden der deutschen Stdpolar-Expedition 1901—1903.— Deutsch Stidpolar-Expedition 18:235-414, figs. 1-68. . 1931. Gammariden und Caprelliden des Magellangebietes, Sudgeorgiens und der Wes- tantarktis —Further Zoological Results of the Swedish Antarctic Expedition 1901-1903, 2(6): 1-290, figs. 1-136, one plate. . 1934. Eine neue Amphipoden-Gattung aus einer belgischen Hohle, nebst Bemerkun- gen lber die Gattung Crangonyx.—Zoologischer Anzeiger 106:215—218, one figure. Shoemaker, C. 1933. Amphipoda from Florida and the West Indies.—American Museum Novitates 598: 1-24, figs. 1-13. . 1964. Seven new amphipods from the West coast of North America with notes on some unusual species.—Proceedings of the United States National Museum 115:391- 430, figs. 1-15. Sivaprakasam, T. E. 1969. Notes on some amphipods from the south east coast of India.— Journal of the Marine Biological Association of India 9:372-383, figs. 14. Sowinsky, V. 1894. Rakoobraznyia Azovskago Moria.—Zapiski Kievskago Obshchestva Es- testvoispytatelei 13:289-405, pls. 8-15. 1898. Vysshiia rakoobraznyia (Malacostraca) Bosfora, po materialam sobrannym d-rom A.A. Ostroumovym ‘‘b’’ 1892 i 93 gg.—Zapiski Kievskago Obshchestva Estest- voispytatelei 15:447-518, pls. 8-13. . 1904. Vvedenie v”’ izuchenie fauny Ponto-Kaspiisko-Aralyskago Morskogo Basseina, razematrivaemoi e’’ tochki erienija eamoetoja telynoi zoo-geograficheskoi provinchie. (Introduction a l’etude de la faune du bassin marin ponto-caspien du point de vue d’une province zoogéographique independante.)—Zapiski Kievskago Obshchestva Estestvois- pytatelei 18:1-487, I—-XIII, pls. 1-4. Stebbing, T. R.R. 1890. The right generic names of some Amphipoda.—Annals and Magazine of Natural History, series 6, 5:192-194. . 1899. Amphipoda from the Copenhagen Museum and other sources. Part II.—Trans- actions of the Linnean Society of London, series 2, Zoology 8:395—432, pls. 30-35. ———. 1906. Amphipoda I. Gammaridea.—Das Tierreich 21: 1-806, figs. 1-127. . 1907. The fauna of brackish ponds at Port Canning, Lower Bengal: Part V.—Definition of a new genus of Amphipoda, and description of the typical species —Records of the Indian Museum 1:159-162, pl. 7. Stock, J. H. 1974. The systematics of certain Ponto-Caspian Gammaridae (Crustacea, Am- phipoda).—Mittheilungen Hamburg Zoologisches Museum Institut 70:75-95, figs. 1-6. , and H. Nijssen. 1965. Eriopisa longiramus n.sp., a new subterranean amphipod from a Red Sea island.—Sea Fisheries Research Station, Haifa, Bulletin 38:28-39, figs. 1-6. VOLUME 92, NUMBER I 165 Stout, V. R. 1912. Studies in Laguna Amphipoda.—First Annual Report of the Laguna Marine Laboratory: 134-149, figs. 79-84. Straskraba, M. 1967. Re-examination of the taxonomic status of Niphargus indicus Chilton (Amphipoda, Gammaridae) and its zoogeographical relations.—Proceedings of the Sym- posium on Crustacea Held at Ernakulam 1:126-132, figs. 1, 2. Tucolesco, I. 1933. Eine neue Amphipodengattung Euxinia.—Zoologischer Anzeiger 103:35- 41, figs. 1-19. Thomson, G. M. 1880. New species of Crustacea from New Zealand.—Annals and Magazine of Natural History, series 5, 6:1-6, pl. 1. Tzvetkova, N. L. 1975a. Pribrezhnye gammaridyi severnykh i dal’nevostochnykh morei SSSR i sopredel’nykh vod rody Gammarus, Marinogammarus, Anisogammarus, Mesogam- marus (Amphipoda, Gammaridae).—Akademiia Nauk SSSR, Zoologicheskii Institut, Izdatel’stvo ‘‘Nauka’’ Leningradskoe Otdelenie, Leningrad, 1975:1—256, figs. 1-89. . 1975b. O novom vide Pleustidae (Amphipoda)-kommensale morskogo ezha s Koman- dorskikh Ostrovov.—Zoologicheskii Zhurnal 54:121—124, one unnumbered figure. , and V. A. Kudrjaschov. 1975. Novyi rod amfipod semeistva Calliopiidae iz severnoi chasti Tichogo Okeana.—Biologija Morja 4:13-23, figs. 1-5. Walker, A. O. 1906. Preliminary descriptions of new species of Amphipoda from the ‘Dis- covery’ Antarctic Expedition, 1902—1904.—Annals and Magazine of Natural History, series 7, 18:13-18. White, A. 1847. List of the specimens of Crustacea in the collection of the British Museum:i— vili, 1-143. London; E. Newman. Wrzesniowsky, A. 1890. Uber drei unterirdische Gammariden.—Zeitschrift fiir Wissenschaft- liche Zoologie 50:600-—724, pls. 27-32. Zimmerman, R. J., and J. L. Barnard. 1977. A new genus of primitive marine hadziid (Am- phipoda) from Bimini and Puerto Rico.—Proceedings of the Biological Society of Wash- ington 89:565—580, figs. 1-5. (GSK) Biological Institute, 81000, Titograd, Yugoslavia; (JLB) Smithson- ian Institution, Washington, D.C. 20560. PROC. BIOL. SOC. WASH. 92(1), 1979, pp. 166-175 THREE NEW SPECIES OF INDO-WEST PACIFIC LIZARDFISH (SYNODONTIDAE) Barry C. Russell and Roger F. Cressey Abstract.—Three new species of Synodus (S. doaki n. sp., S. jaculum n. sp., and §. rubromarmoratus n. sp.) are described from the Indo-West Pacific. The 3 species are distinguished from the known Indo-West Pacific Synodus by the following combination of characters: Synodus doaki with no pigmented area at upper distal corner of operculum, 55-58 vertebrae, 11-12 peritoneal spots, 312 scales above the lateral line, and anterior palatine teeth longest and forming a discrete group; Synodus jaculum with a patch of dense pigment at the base of the caudal fin, 512 scales above the lateral line, 59-62 vertebrae, and the anterior palatine teeth longest and forming a discrete group; Synodus rubromarmoratus with no pigmented area at upper distal corner of operculum, anal-fin base shorter than dorsal-fin base, pos- terior pelvic process wide, 52-56 vertebrae, 12 peritoneal spots, and the anterior palatine teeth not longest and not forming a discrete group. Recent collecting in eastern Australia-New Zealand by Russell and a comprehensive revision (to be published separately) of Indo-West Pacific Synodus by Cressey brought to light the 3 new species of lizardfish de- scribed here. These are among the 18 Indo-West Pacific species now con- sidered valid (Cressey and Randall, in press; Cressey, m.s. in preparation). Type material has been deposited in the following institutions: Australian Museum, Sydney (AMS); Bernice P. Bishop Museum, Honolulu (BPBM); National Museum of New Zealand, Wellington (NMNZ); U.S. National Museum of Natural History, Smithsonian Institution (USNM). Measurements, counts and terminology used follow those of Schultz (1953: p xxi) and Hubbs and Lagler (1958: pp. 19-26). All measurements were made with vernier calipers to the nearest tenth millimeter. Measure- ments and counts for the holotype are given first. Those for the paratypes, where different from the holotype, appear in parentheses. The last dorsal- and anal-fin rays are split, connected at their base. Synodus doaki, new species (Fig. 1) Synodus sp. Doak, 1972: p. 18, plates 6, 7, (Poor Knights Island, New Zea- land). Synodus sp. Allen et al., 1976: p. 380. (Lord Howe Island). VOLUME 92, NUMBER I 167 Fig. 1. Synodus doaki n. sp. Paratype NMNZ 6168, 171.4 mm SL, Poor Knights Island. Holotype.—NMNZ 5676, 159 mm SL, Nursery Bay, Poor Knights Islands (35°30'S, 174°44’E), New Zealand, 19 m, B. C. Russell, 31 November 1970. Paratypes.—NMNZ 5677, 96.5 mm SL, same data as holotype. NMNZ 5678, 140 mm SL, The Canyon, Poor Knights Islands, 30 m, A. M. Ayling, 17 June 1971. NMNZ 6168, 3, 178-209 mm SL, Nursery Cove, Poor Knights Islands, New Zealand, 19 m, B. C. Russell, 4 March 1975. AMS 1.15338, 229 mm SL, Kingston, Norfolk Island (29°04’S, 167°57'E), 9 m, B. and L. Marsh, 4 March 1961. AMS I.18351-001, 85 mm SL, One Tree Island (23°30'S, 152°05'E), Great Barrier Reef, 30 m, B. C. Russell, 23 September, 1974. AMS I.18773-001, 92 mm SL, Phillip Island (29°07’'S, 167°56’E), Nor- folk Island, 15 m, B. C. Russell, 20 September 1975. BPBM 14655, 151.5 mm SL, Observatory Rock, Balls Pyramid (31°45’S, 159°14’E), Lord Howe Island, 31 m, B. C. Russell, 22 February 1973. BPBM 21057, 244 mm SL, Maro Reef (25°25'N, 170°35'W), Leeward Islands, Hawaii, 100 fms, Town- send Cromwell Cr. 77-02, D. Davis, 21 June 1977. USNM 218793, 2, 91- 92.5 mm SL, coast of Kenya (2°42’'S, 40°53’E), W. Indian Ocean, 140 m, Anton Bruun Cruise 8, Station 420A, 6 November 1964. Diagnosis and comparisons.—A species of Synodus with the following combination of characters: dorsal-fin rays (branched and unbranched) 13- 15 (usually 14); anal-fin rays 8-9 (usually 8); pored lateral-line scales 56-58 (usually 56); transverse scale rows 32/7; vertebrae 55-58; combined dorsal and anal procurrent rays 31-35; anterior palatine teeth longest and in a discrete group; peritoneal spots 11-12; posterior pelvic process wide; no conspicuous spots on posterodorsal operculum. This new species can be separated by its discrete group of long anterior palatine teeth from Synodus indicus (Day), S. similis McCullough, S. kaian- us (Gunther), S$. macrops Tanaka, S. sageneus (Waite), and S. rubromar- moratus n. sp., none of which has an anterior group of longer palatine teeth. The following species have higher vertebral counts than S. doaki (55-58): S. capricernis Cressey and Randall (64-65), S. englemani Schultz (59-62), S. ulae Schultz (62-65), and S. jaculum n. sp. (59-62). Synodus fuscus 168 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Tanaka has a narrow posterior pelvic process. Synodus variegatus (La- cepede) has 5% scales above the lateral line. Synodus hoshinonis Tanaka has conspicuous black pigment on the posterodorsal corner of the opercu- lum. Synodus binotatus Schultz has lower vertebral counts (51-55) and fewer peritoneal spots (0-3). Description.—Dorsal-fin rays i, 12 (ii, 1-1, 14); anal-fin rays 9 (8-9); pec- toral-fin rays 12 (12-13); pelvic-fin rays 8; procurrent rays 31-35, 17-19 dorsal, 14-16 ventral; lateral line continuous, 56 (56-58) pored scales; scale rows above lateral line from dorsal origin 312; scale rows below lateral line to midventral line 7; predorsal scales 15 (14-17); vertebrae 55-58. Proportions in standard length: greatest body depth 6.8 (5.7-8.0); head length 3.2 (3.0-3.7); snout length 13.8 (12.9-15.6); snout to dorsal origin 2.4 (2.3-2.5); dorsal origin to adipose origin 2.3 (2.2—2.5); length of dorsal-fin base 5.1 (4.7-6.1); length of anal-fin base 9.9 (10.0-11.6). Proportions in head length: snout length 4.3 (3.8-4.3); maxilla 1.7 (1.4-1.6); horizontal measurement of bony orbit 6.3 (4.3-6.3); least width of bony interorbital 11.1 (8.1-14.0); postorbital length of head 1.7 (1.6—1.9); longest dorsal ray 2.0 (1.8-2.5); length of pectoral fin 2.6 (2.2—2.8); length of pelvic fin 1.4 (0.9— 15) Body tubular, head somewhat depressed, caudal region a little com- pressed. Large cycloid scales on body, extending onto cheeks and oper- culum; 5-6 rows of cheek scales, post-oral portion of cheeks scaly. Snout sharply pointed, longer than broad; the anterior nostril on each side bearing a conspicuous leaflike flap on its posterior margin extending well beyond margin of nares when depressed anteriorly. Adipose eyelid narrow. Inter- obital space concave, occipital region bony, weakly rugose above and be- hind eyes with 3 low bony ridges radiating from behind each eye, these more arborescent and distinct in larger specimens. A single row of for- wardly directed teeth in each jaw and 1 or 2 rows of smaller teeth set below the level of lips. Palatine teeth in an elongate V-shaped pad, teeth pointing backwardly, those in front largest and in a discrete group. Lingual teeth well developed, those on free end of tongue largest and slightly re- curved, in 5—6 rows, teeth of inner 1 or 2 rows smaller. Teeth in jaws, palatines, and tongue caniniform, larger teeth with arrow-shaped tips. Pec- toral fins short, reaching a line from base of pelvic fins to origin of dorsal fin. Outer pelvic ray unbranched and short, the fifth branched ray (sixth ray) longest, reaching just beyond vertical from posterior base of dorsal fin. Posterior bony processes of pelvic girdle broad. Peritoneal color pale whit- ish, 11-12 small black spots along each side of body wall. Color.—Fresh color as follows: Body pale with 8 undulated reddish bars extending from middorsal line almost to midventral line; the first and alter- nate bars paler. Three lengthwise reddish streaks along back and upper sides, one just below middorsal line, one between middorsal line and lateral VOLUME 92, NUMBER 1 169 line, one just above lateral line. A series of reddish blotches along lower part of the sides between first and seventh vertical bars. Head reddish, cheeks and operculum pale, marbled with red. Lips broadly barred with red, a pair of red spots at tip of snout. Eye marked with red, pupil bright red. All fins marked with red transverse bands, adipose fin red, basal portion whitish. An excellent color photograph of this fish is provided by Doak (1972: plate 6). Colors are faded almost completely in alcohol. Distribution and habitat.—Type specimens collected at the Poor Knights Islands in northern New Zealand, where they are locally common at depths of 19-30 m. Specimens collected at Norfolk Island, Lord Howe Island, and at One Tree Island, Great Barrier Reef were taken in water 9-35 m. The spec- imens from Hawaii and the Indian Ocean were trawled in depths greater than 140 m. Synodus doaki inhabits sandy bottoms close to rock or coral outcrops. In New Zealand they frequently were seen in pairs (Russell, pers. observation). The present disparate distribution of this species (Hawaii, E. Australia— northern New Zealand, W. Indian Ocean) suggests a wide distribution, and additional collecting at intervening localities in deeper water undoubtedly will produce additional specimens. Etymology.—Named doaki after Wade Doak, pioneer New Zealand diver and underwater naturalist, who first discovered this fish at the Poor Knights Islands. Synodus jaculum, new species (Fig. 2) Holotype.—AMS 1.19470-005, 62.5 mm SL, Granite Bluff, Lizard Island (14°40’S, 145°27’E), Great Barrier Reef, 16 m, B. C. Russell, 20 November 1975. Paratypes.—AMS 1.17262-053, 77.4 mm SL, Manubada Island (9°30’S, 147°0S'E), Port Moresby, New Guinea, 1.5—3 m, B. Goldman, 24 June 1970. AMS I.17503-014 (2), 55-73.8 mm SL, Blanche Bay (4°18’S, 152°11’E), near Rabaul, New Britain, 4 m, G. R. Allen and J. E. Randall, 7 August 1973. AMS 1.18303-001, 59 mm SL, off Clovelly Pool (33°55’S, 151°17'E), New South Wales, Australia, 12 m, R. Kuiter, 10 February 1975. AMS I.18340- 001, 91 mm SL, One Tree Island (23°30’S, 152°05’E), Great Barrier Reef, 30 m, B. C. Russell, 15 September 1974. AMS I.18340-002 (2), 85.5-91 mm SL, One Tree Island, Great Barrier Reef, 33 m, B. C. Russell, 18 September 1974. AMS 1I.19222-001 (4), 65.3-78.5 mm SL, Sand Cay north of Lizard Island (14°39’S, 145°27'E), Great Barrier Reef, 21 m, B. C. Russell, 1 No- vember 1974. AMS I.19473-054 (2), 59-82 mm SL, south end Coconut 170 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Lern hs, Pe ' rae Fig. 2. Synodus jaculum n. sp. Paratype AMS I.18303-001, 59 mm SL, off Clovelly Pool, New South Wales, Australia. Beach, Lizard Island (14°40’S, 145°27'E), Great Barrier Reef, 2-7 m, AMS party, 24 November 1975. AMS I.19696-004, 123 mm SL, South Solitary Island (30°13’S, 152°19’'E), New South Wales, Australia, 20 m, B. C. Rus- sell, 27 September 1976. USNM 217673, 103.5 mm SL, Pacific Macclesfield Bank (16°3.7'N, 114°43.5’E to 114°40’E), South China Sea 82-84 m, trawled ‘‘Cape St. Mary’’ cruise 3/64, W. L. Chan, 21 June 1964. USNM 217748, 93 mm SL, Pacific Macclesfield Bank (15°33.2’N, 113°56’E to 15°35.5'N, 113°54.5’E), South China Sea, 81-88 m, trawled ‘‘Cape St. Mary’’ cruise 3/ 64, W. L. Chan, 18 June 1964. USNM 217782, 91 mm SL, Pacific Mac- clesfield Bank (16°04.2'’N, 114°41.8’E to 16°04’N, 114°39’E), South China Sea, 81 m, trawled “‘Cape St. Mary’’ cruise 3/64, W. L. Chan, 14 June 1964. USNM 217621 (2), 106-114.3 mm SL, Gulf of Manner (8°39’N, 79°37'E), Ceylon, 6-10 fms, T. Roberts, 3 April 1970. USNM 217640 82.8 mm SL, Comores Islands (12°53’S, 45°16’E), R. V. Anton Bruun, Cruise 9, 26 November 1964. USNM 217794, 79.1 mm SL, Christmas Island (1°56’N, 157°29'W), Line Islands, 3-4 m, R. Bolin, R. V. Te Vega, Cruise 8, 22 July 1965. Diagnosis and comparisons.—A species of Synodus with the following combination of characters: dorsal-fin rays (branched and unbranched) 11- 13 (usually 12); anal-fin rays 8-9 (usually 8); pored lateral-line scales 59-62 (usually 60); transverse scale rows 5%-614/10-11; vertebrae 59-62 (usually 60); nasal flap small, short and triangular; anterior palatine teeth longest and in a discrete group; peritoneal spots 11-13; posterior pelvic process wide; base of caudal fin and peduncle black (colors in alcohol faded, but black on peduncle usually persists). The conspicuous black pigment spot on the caudal peduncle of S$. jaculum distinguishes it from all known species, but the preserved specimens from the South China Sea are much paler than the other material studied, and the black peduncular spot is not present. Synodus jaculum n. sp. can be VOLUME 92, NUMBER 1 171 separated by its discrete anterior group of palatine teeth from S. indicus, S. similis, S. kaianus, S. macrops, S. sageneus, and §. rubromarmoratus in which the teeth are approximately equal in size and do not form a discrete group. It can be separated by its higher vertebral count (59-62) from S. binotatus (51-55), S. fuscus (53-56) and §. hoshinonis (54—56). It can be separated from S$. doaki by its higher number of scales above the lateral line (5.5 vs. 3.5). It differs from S. capricornis by its lower vertebral count (59- 62 vs. 64-65), from S. ulae by its smaller nose flap at the anterior nares and its generally lower vertebral count (59-62 vs. 62-65), from S. englemani by its higher peritoneal spot count (11-13 vs. 7-10), and from S. variegatus by its generally higher vertebral count of jaculum (59-62 vs. 55—60) and its short nose flap (long in variegatus). Description.—Dorsal-fin rays 11, 10 (i, 10-ii, 10); anal-fin rays 8 (8-9); pectoral-fin rays 13 (12-13); pelvic fin rays 8; procurrent rays 28-33, 15-18 dorsal, 13-15 ventral; lateral line continuous, 60 (59-62) pored scales; scale rows above lateral line from dorsal origin 542-6; scales below lateral line to midventral line 10 (10-11); predorsal scales 20 (18-22); vertebrae 59-62. Proportions in standard length: greatest body length 6.9 (5.7-8.1); head length 3.0 (2.9-3.5); snout length 13.9 (13.0-16.9); snout to dorsal origin 2.1 (2.3-2.5); dorsal origin to adipose origin 2.4 (2.3—2.5); length of dorsal-fin base 6.9 (5.6—7.6); length of anal-fin base 10.4 (11.2—18.5). Proportions in head length: snout length 4.7 (4.3-5.1); maxilla 1.4 (1.3-1.7); horizontal measurement of bony orbit 4.7 (4.7-6.0); least width of bony interorbital 8.4 (6.4—11.2); postorbital length of head 1.6 (1.5—1.7); longest dorsal ray 2.1 (1.8-2.8); length of pectoral fin 3.2 (2.6—-3.7); length of pelvic fin 1.3 (1.3- 1.6). Body tubular, head somewhat depressed, caudal region a little com- pressed. Large cycloid scales on body, extending onto cheeks and oper- culum; 4—7 rows of cheek scales, post-oral portion of cheeks naked. Snout pointed, about as long as broad; anterior nostrils on each side bearing a short, triangular flap on its posterior margin, not extending beyond margin of nares when depressed anteriorly. Eye large, adipose eyelid narrow. In- terorbital space concave, occipital region bony, smooth above and behind eyes with 3 bony ridges radiating from behind each eye. A single row of forwardly directed teeth in each jaw 1 or 2 rows of smaller teeth set below level of lips. Palatine teeth in an elongate V-shaped pad, teeth back- wardly pointing, those in front largest and in a discrete group. Lingual teeth well developed, those on free end of tongue largest and slightly recurved, in 4-5 rows, inner 1 or 2 rows of teeth smaller. Teeth in jaws, palatines and tongue caniniform, larger teeth with arrow-shaped tips, particularly in bigger specimens. Pectoral fins short, just reaching to a line from base of pelvic fins to origin of dorsal fin. Outer pelvic ray unbranched and short, the fifth branched ray (sixth ray) longest, reaching to a line drawn vertically 172 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON from posterior base of dorsal fin. Posterior bony processes of pelvic girdle broad. Peritoneal color pale whitish with 11-13 black spots along each side of body wall. Color.—(From color transparencies). Body mottled with red. Nine red- dish undulated bars extending from middorsal line almost to midventral line, each bar broader on the dorsum, the second and every alternate bar lighter and narrower; bars edged with pale blue on dorsum, interspaces pink on dorsum, whitish on upper part of sides; lower part of sides with a reddish bar in interspaces and 2 pairs of white spots, these giving the appearance of 2 broken horizontal white lines along the lower part of the body. Ven- tral surface white. Base of caudal fin and peduncle black, axillary scales white. Head pinkish, cheeks and preopercle whitish, opercle mottled with red. Lips barred with white, 2 blackish streaks extending from anterior nostrils to tip of snout. Eyes reddish, pupils dark, edged with white. Dorsal fin with 3-4 faint transverse dark bands, other fins transparent. Color in alcohol faded almost completely, but the dark mark on the pe- duncle and base of the caudal fin usually persists. Distribution and geographic variation.—Specimens have been collected from Comores Islands, Ceylon, eastern Australia (New South Wales to the northern Great Barrier Reef), New Guinea, New Britain, South China Sea, and the Line Islands. This species occurs in shallow water as well as mod- erate depths (to 88 m) and is probably widespread throughout the Indo-West Pacific region. The higher number of transverse scales above the lateral line (6/2) were on specimens at the periphery of its range, Comores and Christ- mas Island (Pacific). Etymology.—Named jaculum (neuter noun in opposition) from the Latin word for javelin, in reference to the peculiar behavior of this species of launching itself off the bottom and swimming in midwater for prolonged periods. Synodus rubromarmoratus, new species (Fig. 3) Holotype.—AMS 1.19450-027, 75.9 mm SL, Mrs. Watson’s Bay, Lizard Island (14°40'S, 145°27'E), Great Barrier Reef, 15 m, AMS party, 10 No- vember 1975. Paratypes.—AMS 1.19450-024 (10), 50.2-72.0 mm SL, same data as ho- lotype. USNM 218792 (2), 70.3-71.2 mm SL, same data as holotype. Diagnosis and comparisons.—A species of Synodus with the following combination of characters: dorsal-fin rays (branched and unbranched) 10- 12 (usually 11); anal-fin rays 9; pored lateral-line scales 54—55 (usually 54); transverse scale rows 3!4/6; vertebrae 52-55; combined dorsal and ventral procurrent rays 24-29; anterior palatine teeth not longer than posterior and VOLUME 92, NUMBER 1 173 Fig. 3. Synodus rubromarmoratus n. sp. Paratype USNM 218792, 71.2 mm SL, Mrs. Wat- son’s Bay, Lizard Island, Great Barrier Reef. not in a discrete group; pertioneal spots 12; peritoneum pale; anal-fin base shorter than dorsal-fin base; nose flap on anterior nares long and broad. The anterior palatine teeth of S. rubromarmoratus are not longer than those that follow nor do they constitute a discrete group. This character separates them from the following species in which the anterior palatine teeth are longer and in a discrete group: S. binotatus, S. capricornis, S. doaki, S. englemani, S. fuscus, S. hoshinonis, S. jaculum, S. ulae, and S. variegatus. The new species can be separated by its pale peritoneum from S. kaianus and S. macrops (black in the latter 2 species). The lower ver- tebral count of S. rubromarmoratus (52-55) separates it from S. similis (58- 58). In S. sageneus the base of the anal fin is longer than the base of its dorsal fin (shorter in rubromarmoratus). It can be separated from S. indicus by its long, broad, blunt-tipped, nose flap (flap of indicus is long but triangular and sharply pointed) and the higher number of peritoneal spots of (9-11 vs. 12 in indicus). Description.—Dorsal-fin rays i, 9 (ii, 8-ii, 10); anal-fin rays 9; pectoral- fin rays 12 (11-12); pelvic-fin rays 8; procurrent rays 24-29, 13-16 dorsal, 11-14 ventral; lateral line continuous, 54 (54-55) pored scales; scale rows above lateral line from dorsal origin 342; scale rows below lateral line to midventral line 6; predorsal scales 16 (14-16); vertebrae 54 (52-55). Proportions in standard length: greatest body depth 7.1 (7.8—10.0); head length 3.7 (3.7-4.0); snout length 15.8 (15.6—-19.6); snout to dorsal origin 2.3 (2.2—2.4); dorsal origin to adipose origin 2.5 (2.4—2.6); length of dorsal-fin base 8.0 (6.8-8.0); length of anal-fin base 10.0 (9.4-12.2). Proportions in head length: snout length 4.3 (4.1-4.9); maxilla 1.5 (1.3—1.6); horizontal measurement of bony orbit 4.3 (3.6-4.3); least width of bony interorbital 14.8 (11.0-17.5); postorbital length of head 1.7 (1.7—2.1); longest dorsal ray 1.9 (1.5—2.1); length of pectoral fin 2.4 (2.2—2.4); length of pelvic fin 1.3 (1.2- 1.4); measured from tip of snout to rear edge of fleshy operculum. 174 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Body tubular, head somewhat depressed, caudal region a little com- pressed. Large cycloid scales on body, extending onto cheeks and opercule; 5 rows of large cheek scales entirely covering cheeks. Snout pointed, broad- er than long; anterior nostrils on each side bearing a long leaflike flap on its posterior margin extending well beyond edge of nares when depressed an- teriorly. Adipose eyelid narrow. Interorbital space very narrow and con- cave, occipital region bony, smooth, with 3 bony ridges radiating from behind each eye. A single row of forwardly directed teeth in each jaw and 1 or 2 rows of smaller teeth set below level of lips. Palatine teeth in an elongate V-shaped pad, in 2 rows, teeth backwardly pointing, those in front not longer than others. Lingual teeth large and slightly recurved, in 3 rows. Teeth in jaws, palatines and tongue caniniform, larger teeth with arrow- shaped tips. Pectoral fins short, not reaching to a line from base of pelvic fin to origin of dorsal fin. Outer pelvic ray unbranched and short, the fifth branched ray longest, reaching beyond a vertical from posterior base of the dorsal fin. Posterior bony processes of pelvic girdle short, broad. Peritoneal color pale, whitish with 11-12 black spots along each side of body wall. Color.—(From color transparencies). Body mottled with red. Five broad undulated bars extend from middorsal line down to lower half of sides, dark reddish on dorsum, paling to reddish brown on sides. Interspaces mottled with pale reddish brown, a large brownish blotch separating 2 pairs of whitish spots in interspaces on lower part of sides, the spots forming a horizontal row along either side of body. Ventral region pale, whitish. Head reddish brown, cheeks and occipital region bright red. Eye reddish, pupil red, edged with yellow. Fins with 4—5 narrow indistinct transverse reddish bands, adipose fin red. Colors in alcohol faded, head and dorsum mottled with grey, 8 greyish diamond-shaped markings on lower sides. Distribution.—Thus far known only from the Great Barrier Reef, Austra- lia. Type specimens were collected at Lizard Island in the northern part of the Barrier Reef, but this species also occurs in the southern part at One Tree Island (Russell, pers. observation) and is probably widespread through- out the region. Occurs in moderately deep water at depths of 15 m or greater. Etymology.—Named rubromarmoratus (adjective) in reference to the characteristic red marbled coloration of the living fish. Acknowledgments We thank the curators of the various museums for loaning to us much of the material reported here. We also thank Robert Gibbs for critically re- viewing the manuscript and offering a number of helpful suggestions. The illustrations were done by Penelope Kay Hollingsworth. VOLUME 92, NUMBER 1 175 Literature Cited Allen, G. R., D. F. Hoese, J. R. Paxton, J. E. Randall, B. C. Russell, W. A. Starck, F. H. Talbot, and G. P. Whitley. 1976. An annotated checklist of the fishes of Lord Howe Island.—Records of the Australian Museum. 30:365-454. Cressey, R. F. m.s. in prep. Revision of Indo-West Pacific Lizardfishes of the genus Synodus (Synodontidae).—Smithsonian Contributions to Zoology. Doak, W. 1972. Fishes of the New Zealand region.—Hodder and Stoughton, Auckland. 132 pp. Hubbs, C. L., and K. F. Lagler. 1958. Fishes of the Great Lakes Region.—Bulletin of the Cranbrook Institute of Science, No. 26:1-218. Schultz, L. P., and collaborators. 1953. Fishes of the Marshall and Marianas Islands. Volume I. Families from Asymmetrontidae through Siganidae.—Bulletin, U.S. National Museum 202(1): 1-685. (BCR) School of Biological Sciences, Macquarie University, North Ryde, N.S.W. 2113, Australia; (RFC) Smithsonian Institution, Washington, D.C. 20560. PROC. BIOL. SOC. WASH. 92(1), 1979, pp. 176-183 A REEVALUATION OF SESARMA BARBIMANUM CANO, 1889 AND S. CRASSIPES CANO, 1889 (CRUSTACEA: DECAPODA: GRAPSIDAE) Lawrence G. Abele Abstract.—The systematic status of 2 species of brachyuran crabs de- scribed by Cano in 1889 is examined. The types of both species were orig- inally deposited in the Naples Museum but no information on their status is available. Sesarma barbimana, type-locality Payta, Peru, is considered a mislabeled specimen of Nanosesarma minutum (De Man, 1887), a wide ranging Indo-West Pacific species. The second species, Sesarma crassipes, type-locality Pernambuco, Brazil, is considered valid and a male from the Caribbean coast of Costa Rica is described and illustrated. G. Cano in 1889 reported on the Brachyura and Anomura collected during the world cruise made by the Vettor Pisani in 1882 to 1885. Collecting stations were made in the Atlantic, Antarctic, Indo-West Pacific, central Pacific, eastern Pacific and Red Sea. He listed 219 species (not 220 as Stated), including 2 species of the brachyuran genus Sesarma which he described as new. These are S. crassipes, type-locality Pernambuco, [Bra- zil} and S. barbimana, type-locality Payta [Peru]. No material of either species was reported on until 1972 when Coelho and Ramos (1972) listed S. crassipes from estuaries in Pernambuco. Sesarma barbimana remains un- reported from Payta or anywhere else, although Tweedie (1950) suggested that the species should be transferred to the genus Nanosesarma Tweedie, 1950. During a revision of the American species of Sesarma now in progress, it became necessary to determine the status of the above 2 species. Search in crustacean collections of the National Museum of Natural His- tory, Washington, D.C., the Museum of Comparative Zoology, Cambridge, the American Museum of Natural History, New York, the Peabody Museum of Natural History, New Haven, and the Allan Hancock Foundation, Los Angeles, revealed but a single specimen that could be attributed to either species. This is a male from the Atlantic coast of Costa Rica identified as S. crassipes by Dr. Fenner A. Chace, Jr., of the National Museum of Nat- ural History. No other material of either species was found in the identified or unidentified collections. Inquiries to the Naples Museum, the site where the types were originally deposited, have been unanswered. Fieldwork on both coasts of Panama also failed to reveal either species. VOLUME 92, NUMBER 1 177 The present report describes and illustrates the single available male of S. crassipes (it was not possible to obtain the Pernambuco specimens) and suggests that the type-specimen of Sesarma barbimana did not actually come from Payta but is a junior subjective synonym of Nanosesarma min- utum (De Man, 1887), a widespread Indo-West Pacific species. The status of Sesarma barbimana Cano, 1889 Sesarma barbimanum was described from a single female specimen with a carapace length of 6 mm and breadth of 7 mm. The type, if adult, is very small for a species of Sesarma. Cano notes 2 characteristics that preclude placement of S. barbimanum in the genus Sesarma: (1) the chelae are coarsely and densely pilose outside and (2) the merus joints of the legs have a broad dilation below which ends in a strong tooth. No American species of Sesarma has these characteristics and, to my knowledge, no other species in the genus Sesarma shares these characteristics. Tweedie (1950) had already recognized this and placed S. barbimanum in the genus Na- nosesarma Tweedie, 1950. There are 10 nominal species in the genus Na- nosesarma and none occurs in American waters. Serene and Soh (1970) recognized 2 subgenera of Nanosesarma, Nanosesarma and Beanium Se- rene and Soh, 1970; barbimanum would be in the former subgenus. Six species have been placed in the subgenus Nanosesarma: minutum De Man, 1887; jousseaumei Nobili, 1905; gordoni Shen, 1935; pontianacensis DeMan, 1895; tweediei Serene, 1967; vestitum Stimpson, 1858, and of these only N. minutum (De Man, 1887), N. pontianacensis (De Man, 1895) and possibly N. vestitum (Stimpson, 1858) are considered valid by Serene and Soh (1970). The description of S$. barbimanum corresponds to that of N. minutum which is a widespread and common species. It should be pointed out that Tweedie (1950) selected S. andersoni De Man, 1887 as the type- species of Nanosesarma Tweedie, 1950. However, Serene and Soh (1970) incorrectly indicated $. minutum De Man, 1887 as the type-species of Na- nosesarma and included S. andersoni in their subgenus Beanium. They indicated S. batavica Moreira, 1903 (=S. barbimana De Man, 1890 a name preoccupied by S. barbimana Cano, 1889) as the type-species of Beanium. Since the diagnosis of Beanium includes the characteristics of S. andersoni, the type-species of Nanosesarma, Beanium would seem to be a subjective junior synonym of Nanosesarma. Rathbun (1910) pointed out that it is unlikely that many of the species listed by Cano from both Panama and Payta actually were collected there. Of the 13 species listed by Cano from Payta, Rathbun notes that 7 are typical Indo-West Pacific species. The possibility thus exists that Payta is also an incorrect locality for the type-specimen of S. barbimanum. | therefore con- sider S. barbimanum Cano, 1889 to be a junior subjective synonym of N. 178 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON minutum (De Man, 1887). The synonymy, based on Seréne and Soh (1970), would then be: Nanosesarma minutum (De Man, 1887) Sesarma minuta De Man, 1887:650. Sesarma barbimanum Cano, 1889 (?-type-locality Payta). S. (Sesarma) jousseaumei Nobili, 1905:411. Sesarma (Sesarma) gordoni Shen, 1935:27, text-fig. 7. Nanosesarma gordoni: Tweedie, 1950:311. The status of Sesarma crassipes Cano, 1889 Cano reported 11 species of decapods from Pernambuco and, in contrast to his material reported from Payta, all but 2 of the 11 species had been previously recorded from Brazil. These were Actumnus Targionii n.sp. and Sesarma crassipes n.sp.; I can find no reference to the former species; therefore the present report deals only with the latter species. It would appear then that we can be reasonably sure that Pernambuco is the correct locality for Sesarma crassipes. The species, S. crassipes, is mentioned by Cano in the original descrip- tion, De Man (1892), Rathbun (1897, 1918), Tesch (1917), and by Coelho and Ramos (1972). Only Cano (1889) and Coelho and Ramos (1972) had any material of this species available. As noted earlier, I was unable to get any information from the Naples Museum and can only assume that the holotype is no longer extant. There are a few specimens in Brazilian museums but these are not available at this time. I have unsuccessfully attempted to collect material of this species in Panama and Jamaica. Therefore, the fol- lowing illustrations and descriptions are based on the single male from Costa Rica in the National Museum of Natural History (USNM113280). Sesarma crassipes Cano, 1889 Figs. 1, 2 Sesarma crassipes Cano, 1889:93 (type-locality, Pernambuco). Ses. crassipes: De Man, 1892:261. Sesarma crassipes: Rathbun, 1897:90. Sesarma (Sesarma s.s.) crassipes: Tesch, 1917:142. Sesarma (Sesarma) crassipes: Rathbun, 1918:294. Sesarma (Sesarma) crassipes: Coelho and Ramos, 1972:204. The carapace is broader than long (cl/cb = 0.89) with indistinct granules, each with a small amount of pubescence, present on the anterior and lateral regions of the carapace; they are sparse medially and posteriorly. The lateral margins diverge slightly posteriorly. The interorbital region is subdivided VOLUME 92, NUMBER 1 179 Fig. 1. Sesarma crassipes. Male from near Tortuquero, Costa Rica (USNM 113280). into 4 distinct lobes; the median sinus is deeper than the submedial pair. The frontal region is about 0.56 of the carapace breadth; it is concave me- dially and oblique with a very small concave region to the lateral margins which flare very slightly. A distinct groove from the lateral margin of the frontal region extends posteriorly on the dorsal surface of the carapace to about the level of the lateral tooth. The posterior orbital margin is thickened, slightly sinuous and extends anteriorly forming a large, acute, outer orbital angle. A distinct lateral tooth is present on a higher level than the outer orbital angle. There are about 9 oblique granular ridges on the lateral surface of the carapace. The eyes are well developed and pigmented. The basal antennular seg- ment is wide and short; the palp is slightly longer than the width of the basal segment. The basal antennal segment is expanded laterally to form part of the medioventral portion of the orbit; ventrally it forms part of Verwey’s groove. There is a subtriangular area on the pterygostomial region delimited dorsally by Verwey’s groove. The entire suborbital region is covered by short hairs. The third maxillipeds are gaping, exposing the inner mouth parts; they have long setae along the medial edge and an oblique row of pubescence on the merus. The male chelipeds are robust. In the present specimen the right is slightly 180 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 2. Sesarma crassipes. Gonopods from male (USNM 113280). A, Left gonopod in anterior view; B, Left gonopod in posterior view; C-E, Right denuded gonopod in anterior, posterior and medial views respectively. larger than the left. The posterior mesial border of the merus is weakly toothed ending in a distinct notch proximal to the distal margin. The anterior mesial border is strongly toothed and expanded distally. Two rows of pu- bescence are present on the mesial surface. The lateral border of the merus is toothed and ends in a notch proximal to the distal margin. The mesial border of the carpus is delimited into an acute angle by a row of tubercles; VOLUME 92, NUMBER 1 181 below this angle are 2 large and about 3 small tubercles; the lateral border is rounded and the entire surface is covered by short rugae. The dorsal surface of the palm is marked by a distinct row of tubercles which extends beyond the distal margin; the lateral surface of the palm.is smooth to very weakly marked by short rows of tubercles; the mesial surface has about 10 large tubercles and 15-20 smaller tubercles that extend to and weakly delimit the ventral border of the palm. There are 8-12 tubercles present on the dorsal surface of the movable finger extending from the proximal margin and ending proximal to the corneous, spooned tip of the finger; ventrally there is a large, subbasal tooth and 2-3 smaller distal ones. The immovable finger is armed with a large basal tooth, a subequal one distally and 2 weaker teeth proximal to the corneous, spoon-shaped tip. The walking legs increase in length in the order: first, fourth, second and third. For the third walking leg (fourth pereopod) the merus is about twice the length of the carpus which in turn is shorter than the propodus; the dactylus is slightly shorter than the carpus. The merus length is about twice the width; transverse rows of granules are present and a large subdistal tooth is on the dorsal margin. The walking legs have a ventral and dorsal row of thick pubescence ex- tending from the dorsal distal part of the carpus to the distal margin of the propodus where it extends as 3 narrow rows to the distal part of the dac- tylus; ventrally the row begins on the distal part of the propodus and con- tinues as 3 narrow rows on the dactylus. The ventral surface of the propodus is armed with about 3-S pairs of irregularly spaced dark-colored spines; on each side of the ventral distal margin there are about 4—-S dark-colored spines. The abdomen is subtriangular in outline; the length and width of the telson are subequal. The endpiece of the male gonopod is short and subrectangular in form; there is a shallow sinus on the distolateral margin. Measurements.—The single available male is sexually mature and has a cb of 25.5 mm and cl of 22.2 mm. Cano (1889) stated that the male holotype has a cb of 22 mm and a c/ of 18 mm. Type.—The male holotype is presumed to be lost. Type-locality.—Pernambuco, Brazil. Distribution.—The species is known from near Tortuquero, Costa Rica, and Pernambuco, Brazil. | Habitat.—The label accompanying the specimen from Costa Rica indi- cates that it was ‘‘dipnetted along shore’’ about 2 miles above the mouth of the Tortuquero River at Leo’s (USNM 113280). Coelho and Ramos (1972) list the species from estuaries at Pernambuco. Remarks.—The rediscovery of S. crassipes results in 8 species of the subgenus Sesarma now being recognized from the western Atlantic. These are: bidentatum Benedict, 1892; cookei Hartnoll, 1971; crassipes Cano, 1889; curacaoense De Man, 1892; jarvisi Rathbun, 1914; reticulatum Say, 182 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 1817; rectum Randall, 1839; verleyi Rathbun, 1914. The following combi- nation of characters will serve to distinguish S. crassipes from the above species: Carapace not convex, regions clearly delimited; strong anterolateral tooth present on a higher level than outer orbital angle; frontal region slightly more than half carapace breadth; movable finger armed dorsally with 8-12 acute tubercles; merus of fourth pereopod with length about twice width. Acknowledgments I thank the staff of the National Museum of Natural History, Washington, D.C., for the loan of material and for assistance during my visit to that institution. Ms. Teresa Ellis provided Figure 1. Support was provided by NSF grant DEB 75-22583. Literature Cited Benedict, James E. 1892. Decapod Crustacea of Kingston Harbour.—Johns Hopkins Univ. Cincalil(Gn) aie Cano, G. 1889. Crostacei Brachiuri ed Anomuri raccolti nel viaggio della ‘‘Vettor Pisani’ intorno al globo.—Bol. Soc. Nat. Napoli, ser. 1, 3:79-105. Coelho, Petronio Alves, and Marilena De Araujo Ramos. 1972. A constituicao e a distribuicao da fauna de decapodos do litoral leste da America do sul entre as latitudes de 5°N 39°S.—Trab. Oceanogr. Univ. Fed. Pe., Recife 13:133-236. Hartnoll, R. G. 1971. Sesarma cookei n. sp., a grapsid crab from Jamaica (Decapoda, Brach- yura).—Crustaceana 20:257-262, pls. I, II. Man, J. G. De. 1887. Ubersicht der Indo-pacifischen Arten der Gattung Sesarma Say nebst -einer Kritik der von W. Hess und E. Nauck in der Jahren 1865 und 1880 beschrieben Decapoden.—Zool. Jahrb. Syst. 2:639-689, pl. 1. . 1890. Carcinological studies in the Leyden Museum.—Notes Leyden Mus. 12:49-126. 1892. Carcinological notes in the Leyden Museum. No. 6.—Notes Leyden Mus. 14:225-264, pls. 7-10. . 1895. Bericht uber die von Herrn Schiffscapitan Storm zu Atjeh, an den westlichen Kusten von Malakka, Borneo und Celebes sowie in der Java-See gesammelten Deca- poden und Stomatopoden. Theil 2.—Zool. Jahrb. Syst. 9:75-218. Moreira, C. 1903. Nota appendice as contribuicoes para o conhecimento da fauna brazileira. Crustaceos do Brazil.—Arch. Mus. Nacion., Rio de Janeiro 12:111-117. Nobili, G. 1905. Diagnoses preliminaire de 34 espéces et varietés nouvelles et de 2 genres nouveaux de Decapodes de la Mer Rouge.—Bull. Mus. Natn. Hist. Nat. Paris 11:393- 411, 2 text-figs. Randall, J. W. 1839. Catalogue of the Crustacea brought by Thomas Nuttall and J. K. Town- send, from the west coast of North America and the Sandwich Islands, with descriptions of such species as are apparently new, among which are included several species of different localities, previously existing in the collection of the Academy.—Journ. Acad. Nat. Sci. Philadelphia 8:106-147, 5 pls. Rathbun, M. J. 1897. Synopsis of the American Sesarmae, with description of a new species.— Proc. Biol. Soc. Washington 11:89-92. ———. 1910. The stalk-eyed Crustacea of Peru and the adjacent coast.—Proc. U.S. Nat. Mus. 38:531-620, pls. 36—56. —— we s— VOLUME 92, NUMBER 1 183 . 1914. New genera and species of American brachyrhynchous crabs.—Proc. U.S. Nat. Mus. 47:117-129, 5 text-figs., 10 pls. . 1918. The grapsoid crabs of America.—Bull. U.S. Nat. Mus. 97:i-xxii, 1-461, 161 pls., 172 figs. Say, T. 1817. An account of the Crustacea of the United States—Jour. Acad. Nat. Sci. Philadephia 1:57-80, 97-101, 155-169, 235-253, 313-319, 374-401, 423-441, 445-458, 1 Taf. 1817-1818. Serene, R. 1967. Sur deux espéces nouvelles der Brachyoures (Crustacés, Décapodes) et sur une troisieéme peu connue, recoltées dans la region Malaise.—Bull. Mus. Natn. Hist. Nat. Paris (2) 38:817-827. , and C. L. Soh. 1970. New Indo-Pacific genera allied to Sesarma Say 1817 (Brachyura, Decapoda, Crustacea).—Treubia 27(4):387-416, pls. 1-8. Shen, C. J. 1935. On some new and rare crabs of the families Pinnotheridae, Grapsidae and Ocypodidae.—Chinese Jour. Zool. 1:19-40, text-figs. 1-15. Stimpson, W. 1858. Prodromus descriptionis animalium evertebratorum, quae in Expeditione ad Oceanum Pacificum Septentrionalem, a Republica Federata missa, Cadwaladaro Ringgold et Johanne Rodgers ducibus, observit et descriptsit. Pars V. Crustacea Ocy- podoidea.—Proc. Acad. Nat. Sci. Philadelphia 10:93-110. Tesch, J. J. 1917. Synopsis of the genera Sesarma, Metasesarma, Sarmatium, and Clisto- coeloma, with a key to the determination of the Indo-Pacific species.—Zool. Meded. Leiden 3:127-260, 3 pls. Tweedie, M. W. F. 1950. Notes on grapsoid crabs from the Raffles Museum. I. A new genus and description of a new species of the subfamily Sesarminae.—Bull. Raffles Mus. Singapore 23:310-316, text-fig. 1. Department of Biological Science, Florida State University, Tallahassee, Florida 32306. PROC. BIOL. SOC. WASH. 92(1), 1979, pp. 184-194 A NEW SPECIES OF SPILOCUMA (CUMACEA: BODOTRIIDAE: MANCOCUMINAE) FROM THE GULF OF MEXICO Paul E. Omholt and Richard W. Heard Abstract.—Spilocuma watlingi, the second member of its genus, is de- scribed from the shallow, mesohaline waters of the mouth of Mobile Bay, Alabama. S. watlingi can be distinguished from S. salomani by: (1) the dorsal thoracic keel of ovigerous females, (2) the relative lengths and spi- nation of the thoracic appendages, (3) the reduced spination of the uropodal endopod, and (4) the pigmentation pattern. There have been no published records of cumaceans from the coastal and estuarine waters of Alabama. During a preliminary survey of the Cumacea occurring in Mobile Bay and adjacent areas we collected specimens of a new species referable to Spilocuma Watling, 1977. Watling (1977) created the subfamily Mancocuminae to accommodate Pseudoleptocuma Watling, 1977; Mancocuma Zimmer, 1943; and Spilo- cuma Watling, 1977. The members of this subfamily are characterized by the males having less than 4 pairs of pleopods. Pseudoleptocuma is repre- sented by a single species, P. minor (Calman, 1912), having a male with 3 pairs of well developed pleopods. The genus Mancocuma contains 2 species, M. altera Zimmer, 1943 and M. stellifera Zimmer, 1943, both hav- ing males with 2 pairs of reduced pleopods. Pleopods are absent in the genus Spilocuma, which contains S. salomani Watling, 1977 and the new species described in this report. Specimens used in this study were collected at night (2100-2400 hours) by handtowing a #6 (243 um mesh), 0.5 meter plankton net in 1.0 to 1.5 meters of water. Collections were made 7 October and 14 October 1977 along the southern beaches of the eastern tip and middle of Dauphin Island, Mobile County, Alabama. Total length (T.L.) was measured from the infero- lateral edge of the carapace to the end of pleonite 6. Spilocuma watlingi, new species Material examined.—14¢d 5 (T.L. 1.9-2.7 mm), 15 ovigerous 2 @ (T.L. 3.0-4.0 mm), 27 nonovigerous 9? 2 (T.L. 2.2-3.5 mm); southeastern end of Dauphin Island, Alabama (30°14'21"N, 88°4'42”W); night 7 October 1977; 1.0-1.5 meters. 3 do (T.L. 2.0-2.6 mm), | ovigerous 2 (T.L. 3.4 mm), 3 nonovigerous 2? (T.L. 2.7-3.4 mm); middle of Dauphin Island, Alabama VOLUME 92, NUMBER 1 185 ee / A A,B O.Smm a} C,D 0.2mm persian Beaece ce SSS (760 | Fig. 1. Spilocuma watlingi, new species, ovigerous female: A, Lateral view; B, Dorsal view of carapace and thoracic somites; C, Left antenna 1, dorsal view; D, Left antenna 2, dorsal view; E, Uropod, internal lateral view. ee 186 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON (30°14'42”N, 88°11'36’W); night 14 October 1977; 1.0-1.5 meters. Collec- tions made by handtowing a #6 (243 wm mesh), 0.5 meter plankton net; coll. P. E. Omholt and R. W. Heard. Diagnosis. —Female (ovigerous): Third thoracic somite forming distinct keel. Antenna 1, flagellum nearly equal to peduncle segment 3. Antenna 2, second article without setae or spines. Maxilliped 3, carpus with 7-8 internal spines; propodus with 4 internal spines. Uropod, endopod basal segment with 4-6 spines on inner margin. Male: Dorsal keel absent. Antenna 1, articles 1, 2 and accessory flagellum subequal in length. Uropod, endopod basal segment with 5-6 serrated spines on inner margin. Description.—(Based on 16 adult specimens) Ovigerous female (Fig. 1 A and B). T.L. 3.0-4.0 mm. Carapace, 4 total length, extending a short distance in front of ocular lobes; inferolateral edge smooth with small an- tennal sinus. All thoracic somites visible in dorsal view. All ovigerous fe- males with third thoracic somite forming distinct Keel. Dark chromatophores cover carapace and all thoracic and pleon somites except pleonite 6. Antenna 1 (Fig. 1C): Peduncle articles 2 and 3 subequal and together 74 longer than article 1; article 1 with 2-3 short lateral spines; article 2 ringed with 7-8 strong distal spines; article 3 ringed with 5-7 distal spines. Main flagellum of 2 articles; distal article % basal article and together subequal to peduncle article 2; basal article with 1-3 short distal spines; distal article terminating in 1-3 short and 2 long setae plus 2 segmented aesthetascs. Accessory flagellum uniarticulate, /% length basal article of main flagellum; armed with 3 terminal spines. Antenna 2 (Fig. 1D): Small and of 4 articles. Article 3, 2 length article 4, together 24 combined lengths of articles 1 and 2; armed with 1 small lateral spine. Article 4 with 1 lateral and 4 terminal brush setae. Labium (Fig. 2A): Small, of 2 lateral lobes connected at base; densely ciliated along inner edge and outer part of exterior margin. Mandibles (Fig. 2B): Normal in shape; pars molaris large and for grinding; 5 recurved spines along inner margin between pars incisiva and pars molaris. Pars incisiva of left mandible with 5 small teeth; right mandible with 4 small teeth. Well developed lacinia mobilis on left mandible, rudimentary on right. Maxillula (Fig. 2C): Small and of 2 lobes; outer masticatory lobe with 8- 1] terminal spines; inner lobe shorter, with 5-6 curved terminal spines. Palp on basal part of exterior lobe, turning backward terminating in 2 long setae. Maxilla (Fig. 2D): Basis with 2 masticatory expansions; anterior expan- sion with numerous spines; posterior expansion with 8-9 spines. Biarticulate palp originating near basis and directed anteriorly; both lobes similiarly armed with curved spines. First maxilliped (Fig. 2E): Endopod of 5 articles. Basis greater than com- bined lengths of remaining articles; distal internal angel with 3 rows of VOLUME 92, NUMBER 1 187 Fig. 2. Spilocuma watlingi, new species, ovigerous female: A, Right labium; B, Mandibles, dorsal view; C, Right maxillula; D, Right maxilla; E, Left maxilliped 1; F, Left maxilliped 2; G, Right maxilliped 3, lateral view. 188 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON spines; internal margin with 4-5 spines. Article 2 unarmed and '% article 3. Article 3 with 3 internal rows of spines; distal external surface with 1 plu- mose seta. Article 4 with 4—5 internal brush setae. Article 5, 1% article 4; armed terminally with 1 strong and 2 minor spines. Branchial apparatus with 1 accessory and 4 branchial lobes. Second maxilliped (Fig. 2F): Endopod of 6 articles. Basis slightly smaller than combined lengths of remaining segments; distal internal margin with 3 plumose setae; distal external margin with | plumose seta. Ischium un- armed and !/6 merus. Merus subequal to carpus; distal internal surface with 1 plumose seta. Carpus with 6-8 internal spines. Propodus slightly longer than carpus, with 4—S internal spines; external surface with 1 plumose seta and 2 spines. Dactylus % propodus, terminating in 2 strong curved spines. Coxal segment with 7 setae. Third maxilliped (Fig. 2G): Basis greater than combined lengths of re- maining segments; distal angle not produced; distal lateral margin with 2 long plumose setae and 1-2 small spines; internal margin with 9-10 long plumose setae; longitudinal axis of ventral surface with 4—6 spines; borders of distal half outlined with fine hairs. Ischium 14 merus and lacking spina- tion. Merus with 1 plumose seta on distal external angle and 2—3 plumose setae along internal margin. Carpus subequal to merus; | plumose seta on external margin and 7-8 spines along internal surface. Propodus %4 carpus with | lateral spine and 1 distal external spine; internal margin with 4 spines; distal half with 4 tufts of hairs perpendicular to longitudinal axis. Dactylus greater than 12 propodus; armed mid-laterally with | small spine and ter- minally with 3 small spines and 2 stout recurved spines (1 spine % di- ameter of other); 2 tufts of hairs on distal half, 1 perpendicular and the other parallel to the longitudinal axis. Exopod of 7 segments. First pereopod (Fig. 3A): Basis greater than combined lengths of distal articles; 13-16 plumose setae along internal margin; 1 plumose seta on basal external margin; 7-9 small spines along longitudinal axis; distal margin armed externally with 1 long plumose seta and internally with 2—3 plumose setae and 2 spines. Ischium shortest of all segments; distal internal margin with 0-1 spine. Merus 2 times ischium; armed with 2 spines, 1 on distal external margin and 1 on the internal margin. Carpus % longer than pro- podus; armed internally with 3-4 spines and externally with 1-2 distal spines. Propodus slightly smaller than merus; 2 spines along external margin and 3 spines along internal margin. Dactylus #4 propodus, with 2 strong curved terminal spines and 6-7 subterminal spines. Exopod of 7 segments. Second pereopod (Fig. 3B): Shorter than first pereopod. Basis slightly shorter than combined lengths of remaining articles; 13-15 plumose setae along internal margin; 2 plumose setae on basal external margin; 8-9 simple spines along longitudinal axis of ventral surface; distal margin with 1 internal and | external seta. Merus smaller than carpus or dactylus but larger than VOLUME 92, NUMBER 1 189 A-C 0.2mm > RARUARDAL ~ A (@) \, Oy aco FS Fig. 3. Spilocuma watlingi, new species, ovigerous female: A, Left pereopod 1, lateral | | view; B, Right pereopod 2, lateral view; C, Right pereopod 3, lateral view. a 190 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON FEO Fig. 4. Spilocuma watlingi, new species, ovigerous female: A, Left pereopod 4, lateral view; B, Right pereopod 5, lateral view. propodus; distal internal margin with | strong serrated spine; distal external margin with 1 segmented spine. Carpus subequal to dactylus; 1 strong ser- rated spine and 2 smaller spines on distal internal margin; distal external margin with 2 short segmented spines. Propodus with | strong serrated spine and 2-3 simple spines on distal internal surface. Dactylus with 3 strong terminal spines and 5-6 strong subterminal spines. Exopod of 7-8 segments. Third pereopod (Fig. 3C): Shorter (°/,9) than first pereopod. Basis sub- equal to combined lengths of remaining articles; 2-3 plumose setae along basal internal margin; internal margin with 4—5 simple spines; external sur- face with 4—S plumose setae. Ischium % merus; distal margin ringed with 5-6 setae (3 setae extending to distal margin of carpus). Merus subequal to carpus; external margin with 4-5 spines. Carpus with 1-2 spines on mid- lateral surface; distal margin with 3 long setae extending past dactylus. Propodus 4% carpus; distal margin with 1 long seta. Dactylus 24 propodus with | strong curved terminal spine and 1-2 subterminal spines. Exopod of 7 segments. Fourth pereopod (Fig. 4A): 4/s first pereopod. Basis 4/5; combined lengths of remaining segments; external margin with 7 plumose setae; distal internal margin with | segmented and 2 simple spines. Ischium subequal to propodus; distal surface ringed with 6 segmented setae. Merus 2 times is- VOLUME 92, NUMBER | 191 Fig. 5. Spilocuma watlingi, new species, adult male: A, Lateral view; B, Right antenna 1, dorsal view; C, Right antenna 2, lateral view; D, Uropod, internal lateral view. chium; internal margin with 4—S short segmented setae. Carpus longest of the S distal segments; external margin with 2—3 short segmented setae; distal margin with 3 long segmented setae extending past dactylus. Propodus with 1 short spine on longitudinal axis of ventral surface; | long segmented seta on distal margin equal to those on carpus. Dactylus 74 propodus; armed with 1 strong recurved terminal spine and 1-2 subterminal spines. Rudi- mentary exopod indistinctly jointed terminating in 2 branching setae. 192 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fifth peropod (Fig. 4B): 3/5 first pereopod. Basis 4 times ischium; longitudinal axis of ventral surface with 4-5 plumose setae; distal surface with 2 segmented setae. Ischium ringed with 3-4 distal setae. Merus smaller than carpus; distal surface with 3-4 setae. Carpus with 2 setae on distal surface extending past dactylus; internal surface with 2-3 setae. Propodus 14 merus; distal surface with 1 long seta equal to those on distal surface of carpus, and 1 small spine. Dactylus #4 propodus, terminating in | strong stout spine. No exopod present. Uropod (Fig. 1E): Peduncle with 3-6 strong simple spines along inner margin. Endopod of 2 articles, distal article % basal article; basal article with 4-6 stout serrated spines along inner margin; distal article with | ser- rated lateral spine and 2 terminal spines. Exopod with 3 lateral and 3 ter- minal strong spines. Male (based on 17 adult specimens): T.L. 1.9-2.7 mm. Body and pig- mentation pattern similar in appearance to adult female except thoracic keel absent. (Fig. 5A). Antenna 1 (Fig. 5B): Peduncle of 3 articles; article 1 with 2—3 short lateral spines; article 2 ringed with 12-13 stout distal spines; article 3 longer than articles | or 2 with 6 stout distal spines. Flagellum of 3 articles; article 1 subequal to article 2; article 3, % article 2. Article 1 with 2 aesthetascs and 1-3 short spines; article 2 with 1-2 short distal spines and | long distal seta; article 3 with 2 aesthetascs and 2—3 long setae. Accessory flagellum uniar- ticulate and subequal to article 1 of flagellum; armed with 3-4 terminal setae. Antenna 2 (Fig. SC): Modified for grasping. Peduncle of 3 articles; articles 1 and 2 without spines; article 3 longer than combined lengths of articles 1 and 2, with 2 basal spines, 5-8 distal setae and 10-12 small mounds with single flagellum along internal axis. Flagellum with 15 articles, each bearing flattened granulated adhesive pads on internal side, except last 4 articles. Article 1 of main flagellum bears 1 pad, articles 2-11 bear 3 pads each; articles 12 and 14 with 2 fine spines and no pads; article 13 without pads or spines; last article (15) of flagellum terminating in 3 fine spines. Uropod (Fig. 5D): Peduncle inner margin with 5-6 strong spines. Endopod basal article with 5-6 stout pronged spines on inner margin; distal article with | lateral pronged spine and 2 terminal strong simple spines. Exopod with 1 lateral and 3 terminal spines. Holotype.—Ovigerous female, USNM No. 171296; T.L. 3.1 mm. Paratypes.—7 adult males and 8 females, USNM No. 171297; 366 T.L. 2.0-2.4 mm, 2 2 T.L. 3.1-4.0 mm. Type-locality.—Southeastern end of Dauphin Island, Alabama, (30°14’21°N, 88°4'42°W). Depth 1.0-1.5 m. Habitat.—In the shallow, mesohaline waters of protected beaches adja- cent to mouths of bays or estuaries. VOLUME 92, NUMBER 1 193 Table 1. Selected morphological comparison of S. watlingi with S. salomani. Feature S. watlingi S. salomani pigmentation (d + @) Female Antenna | main flagellum Antenna 2 second article Maxilliped 3 carpus propodus Pereopod | propodus Uropod endopod basal distal exopod Male Antenna 1 main flagellum Uropod peduncle endopod basal distal exopod absent from pleonite 6 subequal to peduncle 3 no spines or setae 7-8 internal spines 4 internal spines 3 internal spines 4-6 serrated spines 1 lateral and 2 terminal spines 3 lateral and 3 terminal spines articles 1, 2 and accessory flagellum subequal in length 5-6 spines 5-6 serrated spines 1 lateral and 2 terminal spines 1 lateral and 3 terminal spines absent from thoracic somite 5 and pleonites 3-4 Y% peduncle 3 plumose setae 10-12 internal spines 10 internal spines 6 internal spines 9-12 spines 3 lateral and 1 terminal spines 5 lateral and 1 terminal spines article 2 > accessory > article 1 6-7 spines 12-15 serrated spines 1 lateral and 4 terminal spines 2 lateral and 2 terminal spines Etymology.—This species is named in honor of Dr. Les Watling, Ira C. Darling Center, University of Maine. Discussion With the present description of Spilocuma watlingi the genus Spilocuma now contains 2 species, both recorded from the shallow waters of the eastern Gulf of Mexico. Morphologically S. watlingi can be distinguished 194 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON from S. salomani by: (1) the presence of a dorsal Keel on the third thoracic somite of ovigerous females, (2) having fewer spines on the inner margin of both segments of the uropodal endopod, (3) the spination and relative lengths of the thoracic appendages, and (4) the pigmentation pattern. These and other differences between the 2 species are compared in Table I. Spilocuma watlingi and S. salomani also appear to have different habitat preferences. We have examined large numbers of S. salomani from the beaches of Panama City and Pensacola, Florida, and Dauphin Island, Ala- bama. In all instances these samples came from high energy beaches. Sam- ples collected from the protected low energy beaches at the southeastern tip of Dauphin Island contained S. watlingi but no S. salomani. Collections made on high energy beaches at the middle of Dauphin Island, approxi- mately 4 miles west of the eastern tip, contained large numbers of S. salomani and only a few specimens of S. watlingi. These limited data suggest that S. salomani is restricted to high energy beaches, whereas, S. watlingi generally occurs along low energy beaches under more estuarine conditions. Acknowledgments We wish to thank Michael R. Dardeau and Daniel L. Adkison for their critical review of the manuscript. Contribution No. 26 of the Marine En- vironmental Sciences Consortium, Dauphin Island, Alabama 36528. Literature Cited Watling, L. 1977. Two new genera and a new subfamily of Bodotriidae (Crustacea: Cumacea) from eastern North America. Proc. Biol. Soc. Wash. 89:593-598. (PEO) Department of Biology, University of Alabama in Birmingham, Birmingham, Alabama 35294; (RWH) Gulf Coast Research Laboratory, Ocean Springs, Mississippi 39564. Present address.—(PEO) Dauphin Island Sea Lab., P.O. Box 386, Dauphin Island, Alabama 36528. PROC. BIOL. SOC. WASH. 92(1), 1979, pp. 195-203 CORRECT CITATIONS FOR SOME NORTH AMERICAN BIRD TAXA Richard C. Banks and M. Ralph Browning Abstract.—Correct citations are provided for 24 bird taxa that are incor- rectly cited in current standard references and synonymies. The corrections result from a determination of actual publication dates of a series of papers by Robert Ridgway in late 1873 and early 1874. Robert Ridgway was a prolific writer. He must have worked on several manuscripts simultaneously in the early 1870’s, but not all his manuscripts were published in the order in which they were prepared or submitted. This is particularly true of late 1873 and early 1874, when several works con- taining descriptions of new bird taxa appeared over a short period. Even then there were publication lags, journals were not always published on the cover dates, and dates of publication were not always firmly fixed. Numbers 11 and 12 of Volume 5 of the Bulletin of the Essex Institute, published in Salem, Massachusetts, carried an ornithological paper by R. Ridgway (1873c) and one by Spencer F. Baird and Ridgway (1873). The coauthored paper is a continuation of an article that had appeared in the American Naturalist under the byline of Ridgway (1873a) alone, although Baird contributed descriptions of some birds named therein. Each number of the Bulletin was considered a monthly issue, and Numbers 11 and 12 are for November and December 1873, respectively. In December 1873 (1873b) and January 1874, Ridgway had articles appear in two popular sporting newspapers. Finally, the first two volumes of the long-awaited Birds of North America by Baird, T. M. Brewer, and Ridgway were issued in early 1874. Many bird taxa were first described in one of this group of publica- tions, and some taxa were described as many as three times. We wish to show which of the multiple descriptions was actually the first, and to provide correct citations for those taxa that are incorrectly cited in current standard reference works and synonymies. A few other taxonomists seem to have become aware that the dates usually cited are incorrect and have given proper citations for species of their particular interest, but it seems worthwhile to give here the basis for those actions. Our determination of the first publication of these names does not result in the changing of any names presently in use or involve the designation of type specimens in most instances, but in some cases it affects the usually cited authorship. The U.S. Library of Congress (Smithsonian Deposit) contains a bound Volume 5 of the Bulletin of the Essex Institute, which is essentially the 196 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON proceedings of the Institute for 1873. The last page of each of the 12 numbers of the Bulletin bears a stamped date, which we believe to be the date when the number was received by the Smithsonian Institution. For example, Number 1, for January 1873, bears the date 17 June 1873; Number 8, for August, has the date 30 December 1873. The date on Numbers 9 and 10 is 17 February 1874, and that on Numbers 11 and 12 is 2 May 1874. Numbers 11 and 12 were received at the Museum of Comparative Zoology on 23 March 1874 (W. D. Myers, personal communication). These dates suggest that preparation, printing, and distribution routinely took several months. The fact that Number 12 includes information from an Institute meeting of 16 December 1873 virtually precludes the possibility that this number was printed in that year. Separates of Ridgway’s papers from the Bulletin and other sources are bound into volumes housed in the library of the Smithsonian Institution (Division of Birds). His papers from Numbers 11 and 12 of Volume 5 of the Essex Institute carry a terminal printed note (not found on the numbers bound in the Library of Congress volume), ‘Printed at the Salem Press, Feb., 1874.’ This might be the date of printing of the separates, but un- doubtedly approximates the date of printing of the entire number. The actual date of publication cannot be fixed beyond this month and year, and ac- cording to the rules of the International Commission of Zoological Nomen- clature (1964) it must be considered as being the end of the month, or 28 February 1874. Volumes 1 and 2 (of 3) of Baird, Brewer, and Ridgway (1874) were also published in February 1874 (Zimmer 1926:35). Records compiled and main- tained by Charles W. Richmond in the Smithsonian Institution indicate that these two volumes were available by 14 February 1874, and that date may be fixed as the actual date of publication. Thus, descriptions in Volumes 1 and 2 of Baird, Brewer, and Ridgway (1874) predate and have priority over those in the last two numbers of the Bulletin of the Essex Institute for 1873. Both are predated by papers of Ridgway in Forest and Stream (18 December 1873) and American Sportsman (3 January 1874). In the accounts that follow we give first the name of the taxon as currently used, with the correct author and date. Below that we give full citations for the first description and others, if any, in the series of papers under consid- eration. Comments follow these citations where necessary for clarification. Falco peregrinus pealei Ridgway 1874 Falco communis, var. Pealei Ridgway, in Baird and Ridgway, Bull. Essex Inst. 5(12):201. Dec. 1873 (=Feb. 1874). The corrected date appears in the fifth edition of the American Ornithol- VOLUME 92, NUMBER 1 197 ogists’ Union (1957) check-list, but the erroneous one is given in earlier versions of that list and in the standard references including Deignan (1961). Falco columbarius suckleyi Ridgway 1874 Falco columbarius, var. Suckleyi Ridgway, in Baird and Ridgway, Bull. Essex Inst. 5(12):201. Dec. 1873 (=Feb. 1874). As with F. p. pealei, the date is correct as given by the A.O.U. (1957) but incorrect in other standard references. Dendragapus obscurus fuliginosus (Ridgway) 1873 Canace obscura, var. fuligniosa [sic] Ridgeway [sic], Forest and Stream 1(19):289. 18 Dec. 1873. Clanace]. obscurus var. fuliginosus Ridgway, American Sportsman 3(14):210. 3 Jan. 1874. Canace obscura, var. fuliginosa Ridgway, in Baird and Ridgway, Bull. Essex Inst. 5(12):199. Dec. 1873 (=Feb. 1874). Ridgway apparently intended the paper in the Bulletin of the Essex In- stitute to be the description of this subspecies, and in later works he ob- viously considered it so. Only in that paper were type specimens designated. Nonetheless he had provided both a name, description, and range in the two papers that appeared earlier, neither of which, incidentally, is cited in the extensive synonymy in Ridgway and Friedmann (1946). The misspelling of the subspecific name in the Forest and Stream article is obviously a type setter’s error that was emended by Ridgway in later papers. Similarly, the misspelling of Ridgway’s name as author of the paper is an obvious lapsus. Browning (in press) was the first to provide the citation to the ‘‘original’’ description. Tympanuchus pallidicinctus (Ridgway) 1873 Cupidonia cupido var. pallidicinctus Ridgeway [sic], Forest and Stream 1(19):289. 18 Dec. 1873. Cupidonia cupido var. pallidicincta Ridgway, Amer. Sportsman 3(14):40. 3 Jan. 1874. Cupidonia cupido, var. pallidicincta Ridgway, in Baird and Ridgway, Bull. Essex Inst. 5(12):201. Dec. 1873 (=Feb. 1874). The history of this name is the same as that of Dendragapus o. fuligi- nosus. Tyto alba guatemalae (Ridgway) 1874 Strix flammea, var. Guatemalae Ridgway, in Baird and Ridgway, Bull. Essex Inst. 5(12):200. Dec. 1873 (=Feb. 1874). 198 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Otus asio floridanus (Ridgway) 1874 Scops asio, var. Floridanus Ridgway, in Baird and Ridgway, Bull. Essex Inst. 5(12):200. Dec. 1873 (=Feb. 1874). Cory (1919) cited the third volume of Baird, Brewer and Ridgway (1874) as the original reference, but that volume apparently was not published until December 1874 (Zimmer 1926:35). Otus trichopsis trichopsis (Wagler) 1832 Scops asio, var. enano Baird and Ridgway, Bull. Essex Inst. 12(5):200. Dec. 1873 (=Feb. 1874). This name was published as “‘Scops asio. var. enano Lawrence. Ms.”’ Although most names proposed in this paper by Baird and Ridgway are therein attributed to Ridgway alone, this one must under the rules stand as a jointly proposed name. Ridgway (1914:707) included the name in the syn- onymy of Otus trichopsis as ‘‘Scops asio var. enano ‘Lawrence, MSS.’ Ridgway’’ and (op cit:694) mentioned it under Otus asio mccalli as ‘‘Scops asio, var. enano Lawrence.’’ Moore and Peters (1939:45) placed this name in the synonymy of O. f¢. trichopsis and corrected the authorship but not the date. Strix varia sartorii (Ridgway) 1874 Syrnium nebulosum, var. sartorii Ridgway, in Baird and Ridgway, Bull. Essex Inst. 5(12):200. Dec. 1873 (=Feb. 1874). Perisoreus canadensis obscurus Ridgway 1874 [Perisoreus canadensis] var. obscurus Ridgway, in Baird, Brewer and Ridg- way, Birds N. Amer. 11:298, 302. 14 Feb. 1874. Perisoreus canadensis var. obscurus Ridgway, Bull. Essex Inst. 5(11):194. Nov. 1873 (=Feb. 1874). Perisoreus Canadensis, var. obscurus Ridgway, in Baird and Ridgway, Bull. Essex Inst. 5(12):199. Dec. 1873 (=Feb. 1874). Ridgway apparently intended the December 1873 Essex publication to be the original description, for he footnoted the November use as a manuscript name. However, the description in Baird, Brewer, and Ridgway (1874) must Stand as the first. Perisoreus canadensis capitalis Baird 1874 [Perisoreus canadensis] var. capitalis Baird, in Baird, Brewer, and Ridg- way, Birds N. Amer. 1i:298, pl. 41 fig. 4, 302. 14 Feb. 1874. VOLUME 92, NUMBER 1 199 Perisoreus canadensis, var. capitalis Ridgway, Bull. Essex Inst. 5(11):193. Nov. 1873 (=Feb. 1874). Perisoreus canadensis, var. capitalis Baird, in Baird and Ridgway, Bull. Essex Inst. 5(12):199. Dec. 1873 (=Feb. 1874). Ridgway has been considered the author of the name capitalis by those who have accepted the Essex Institute paper as the first use, even though he intended to credit it to Baird as a manuscript name. Recognizing Baird, Brewer, and Ridgway (1874) as the place of original description does credit the name to Baird, even though Ridgway was responsible for most of the taxonomic work in those volumes. Aphelocoma coerulescens sumichrasti (Ridgway) 1874 C.[yanocitta] californica var. sumichrasti Ridgway, in Baird, Brewer, and Ridgway, Birds N. Amer. 11:283, pl. 40 fig. 2. 14 Feb. 1874. Cyanocitta floridana, var. Sumichrasti Ridgway, in Baird and Ridgway, Bull. Essex Inst. 5(12):199. Dec. 1873 (=Feb. 1874). Aphelocoma ultramarina arizonae (Ridgway) 1874 C.[yanocitta] ultramarina, var. arizonae Ridgway, in Baird, Brewer, and Ridgway, Birds N. Amer. 1i:284, pl. 41 fig. 2, 292. 14 Feb. 1874. Cyanocitta Floridana, var. Sumichrasti Ridgway, in Baird and Ridgway, Bull. Essex Inst. 5(12):199. Dec. 1873 (=Feb. 1874). The bird illustrated in figure 40 of Baird, Brewer, and Ridgway (1874) is captioned with the museum number 42129. The USNM specimen that bears that number is of a different species; the number as printed is obviously an error for 42149, the catalog number of the bird considered the type in the other publication (Baird and Ridgway 1873). Lanius excubitor algeriensis Lesson 1839 Collurio ludovicianus, var. robustus Baird, in Ridgway, Amer. Nat. 7(10):609. 1873. Baird contributed the description of this form to Ridgway’s paper, and Ridgway (1897, 1904) credited the name to Baird (cf. Dendroica dominica, beyond). Hellmayr (1935), however, considered it a Baird manuscript name published by Ridgway. The name has not since, to our knowledge, been used in the American literature. 200 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Coereba flaveola caboti (Baird) 1873 [(Certhiola| Caboti Baird, in Ridgway, Amer. Nat. 7(10):612. 1873. Coereba flaveola newtoni (Baird) 1873 [Certhiola| Newtoni Baird, in Ridgway, Amer. Nat. 7(10):611. 1873. Coereba flaveola barbadensis (Baird) 1873 [Certhiola| Barbadensis Baird, in Ridgway, Amer. Nat. 7(10):612. 1873. Coereba flaveola bartholemica (Sundevall and Sparrmann) 1869 [Certhiola] frontalis Baird, in Ridgway, Amer. Nat. 7(10):612. 1873. Ridgway’s (1873a) article in the American Naturalist incorporated a syn- opsis of the genus Certhiola (=Coereba) written by Baird, in which these four forms were described. Ridgway (1902) properly credited the names to Baird. Hellmayr (1935) noted them as Baird manuscript names attributed to Ridgway. Deignan (1961) provided correct citations for frontalis and bar- badensis, but Lowery and Monroe (in Paynter 1968) considered them to be Ridgway’s names, although not providing a citation for frontalis which is considered a synonym of bartholemica. Dendroica dominica albilora Baird 1873 Dendroica dominica, var. albilora Baird, in Ridgway, Amer. Nat. 7(10):606. 1873. In 1902, Ridgway headed the account of this form ‘‘Dendroica dominica albilora Ridgway’? and in the synonymy ambiguously listed the reference to the original description as ‘‘Dendroica dominica albilora Baird, Ridgway, Am. Nat... .’’ Hellmayr (1935), the A.O.U. (1957), Deignan (1961), and Lowery and Monroe (in Paynter 1968) all consider albilora a Baird manu- script name published by Ridgway. However, the name of this taxon is clearly attributed to Baird in Ridgway’s (1873a) paper, in which he stated that “Included in the paper are some hitherto unpublished descriptions of races of birds by Prof. Baird.’’ This situation is identical to that of Lanius ludovicianus robustus, which Ridgway properly (1904) attributed to Baird. Dolichonyx oryzivorus (Linnaeus) 1758 Dolichonyx oryzivorus, var. albinucha Ridgway, Bull. Essex Inst. 5(11):192. Nov. 1873 (=Feb. 1874). Dolichonyx oryzivorus, var. albinucha Ridgway, in Baird and Ridgway, Bull. Essex Inst. 5(12):198. Dec. 1873 (=Feb. 1874). VOLUME 92, NUMBER 1 201 Hesperiphona vespertina montana Ridgway 1874 H.[esperiphona|] vespertina var. montana Ridgway, in Baird, Brewer, and Ridgway, Birds N. Amer. 1:449, pl. 22 fig. 4. 1874. Hesperiphona vespertina, var. montana Ridgway, Bull. Essex Inst. 5(11):181 (nomen nudum), 189. Nov. 1873 (=Feb. 1874). The volume by Baird, Brewer, and Ridgway (1874) was considered the place of original description of this taxon until Zimmer (1953) suggested that the description in Ridgway (1873c = 1874) had been overlooked. Reestab- lishing the former as the first description (as was done by Howell in Paynter 1968 for the wrong reason) adds support to the argument by Grinnell (1917) and Zimmer (1953) that the specimen illustrated therein (USNM 35150) must be considered the holotype. The long controversy over the type of this taxon (see Deignan 1961) is fraught with errors. Leucosticte australis Ridgway 1874 Leucosticte tephrocotis, var. australis Ridgway, Bull. Essex Inst. 5(11):189. Nov. 1873 (=Feb. 1874). Leucosticte tephrocotis, var. australis Baird and Ridgway, Bull Essex Inst. 5(12):197. Dec. 1873 (=Feb. 1874). As noted, this form was described twice; in both instances the name was taken from a manuscript of J. A. Allen and the name was attributed to him. Ridgway (1901) continued to give Allen as the author of the name. The A.O.U. (1886) Check-list also listed australis as an Allen name and cited only the second description, from page 197. All subsequent editions of the Check-list have attributed the name to Ridgway but have continued to cite the second description of this bird. Hellmayr (1938) cites the reference from page 189 but Howell (in Paynter 1968) mentions only the second description. Ammospiza maritima nigrescens (Ridgway) 1874 Ammodramus maritima, var. nigrescens Ridgway, in Baird and Ridgway, Bull. Essex Inst. 5(12):198. Dec. 1873 (=Feb. 1874). Amphispiza bellii nevadensis (Ridgway) 1874 P.[oospiza] belli var. nevadensis Ridgway, in Baird, Brewer, and Ridgway, Birds N. Amer. 1:590, pl. 26 fig. 9, 594. 14 Feb. 1874. Poospiza belli, var. nevadensis Ridgway, Bull. Essex Inst. 5(11):191. Nov. 1873 (=Feb. 1874). Poospiza Bellii, var. Nevadensis Ridgway, in Baird and Ridgway, Bull. Essex Inst. 5(12):198. Dec. 1873 (=Feb. 1874). 202 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Zonotrichia leucophrys gambelii (Nuttall) 1840 Zonotrichia leucophrys, var. intermedia Ridgway, in Baird and Ridgway, Bull. Essex Inst. 5(12):198. Dec. 1873 (=Feb. 1874). Acknowledgments We thank George E. Watson who commented on the manuscript and aided in determining publication dates. William D. Myers of the Museum of Comparative Zoology, Janet Evans of the Academy of Natural Sci- ences, Philadelphia, and Irene R. Norton of the Essex Institute assisted in determining publication dates. Michael Bogan and Alfred Gardner pro- vided useful ideas. Literature Cited American Ornithologists’ Union. 1886. The code of nomenclature and check-list of North American birds. New York. . 1957. Check-list of North American birds. Edition 5. Baltimore, Maryland. Baird, S. F., T. M. Brewer, and R. Ridgway. 1874. A history of North American birds. Land Birds. 3 Vols.—Boston. Little, Brown and Co. , and R. Ridgway. 1873 (=1874). On some new forms of American birds.—Bull. Essex Inst. 5(12):197—201. Browning, M. R. In press. Type specimens of birds collected in Oregon.—Northwest Sci. Cory, C. B. 1919. Catalogue of birds of the Americas.—Field Mus. Nat. Hist., Zool. Ser., 13, pt. 2:1-607. Deignan, H. G. 1961. Type specimens of birds in the United States National Museum.—U.S. Natl. Mus. Bull. 221. Grinnell, J. 1917. The subspecies of Hesperiphona vespertina.—Condor 14:17-22. Hellmayr, C. E. 1935. Catalogue of birds of the Americas.—Field Mus. Nat. Hist., Zool. Ser., 13, pt. 8:1-541. . 1938. Catalogue of birds of the Americas.—Field Mus. Nat. Hist. Publ., Zool. Ser., 13, pt. 11:1-662. International Commission on Zoological Nomenclature. 1964.—International code of zoological nomenclature. London. Moore, R. T., and J. L. Peters. 1939. The genus Otus of Mexico and Central America.—Auk 56:38—S6. Paynter, R. A., Jr., ed. 1968. Check-list of birds of the world. Vol. 14.—Mus. Comp. Zool., Cambridge, Mass. Ridgway, R. 1873a. On some new forms of American birds.—Am. Nat. 7:602-619. Ridgeway [sic] = Ridgway, R. 1873b. The grouse and quails of North America.—Forest and Stream 1, no. 19:289-290. Ridgway, R. 1873c (=1874). The birds of Colorado.—Bull. Essex Inst. 5(11):174-195. . 1874. The nomenclature of American game birds—Am. Sportsman 3:210-211. 1897. On the status of Lanius robustus Baird as a North American bird.—Auk 14:323. 1901. The birds of North and Middle America. Pt. 1.—U.S. Natl. Mus. Bull. 50. 1902. The birds of North and Middle America. Part 2.—U.S. Natl. Mus. Bull. 50. 1904. The birds of North and Middle America. Part 3.—U.S. Natl. Mus. Bull. 50. 1914. Birds of North and Middle America. Pt. 6.—U.S. Natl. Mus. Bull. 50. LT VOLUME 92, NUMBER 1 203 , and H. Friedmann. 1946. Birds of North and Middle America. Pt. 10.—U.S. Natl. Mus. Bull. 50. Zimmer, J. T. 1926. Catalogue of the Edward E. Ayer ornithological library. Part 1.—Field Mus. Nat. Hist., Zool. Ser., vol. 16. . 1953. The original description of Hesperiphona vespertina montana Ridgway.—Auk 702213, National Fish and Wildlife Laboratory, U.S. Fish and Wildlife Service, National Museum of Natural History, Washington, D.C. 20560. PROC. BIOL. SOC. WASH. 92(1), 1979, pp. 204-207 RANGE EXTENSION OF PENAEOPSIS SERRATA (CRUSTACEA: PENAEOIDEA) TO OFF NEW JERSEY AND RIO GRANDE DO SUL Isabel Pérez Farfante and Boris G. Ivanov Abstract.—Penaeopsis serrata, a wide ranging amphi-Atlantic shrimp, has been reported to range in the western Atlantic from off Cape Lookout, N.C., to Surinam. Herein it is recorded from localities off New Jersey and Rio Grande do Sul, about 800 km northward and 6400 km southeast of its previously known limits. This amphi-Atlantic species was previously known to occur in the western Atlantic from off Cape Lookout, N.C., (Hay and Shore, 1918) through the Gulf of Mexico and the Caribbean Sea south to Surinam (Bullis and Thomp- son, 1965). In the eastern Atlantic, it ranges from south of Cabo San Vicente, Portugal, to Tamzak (“‘Tamxat’’), 17°26'N, 16°02'W, Mauritania (Maurin, 1961, 1968). Like its congeners, all from the Indo-West Pacific, P. serrata Bate, 1881, occurs on upper continental and insular slopes at depths of 183 to 750 m. This is a common shrimp, known by fishermen as ‘‘megalops’’—a name proposed by S. I. Smith (1885) and repeatedly used in the literature. In the eastern Atlantic, although quite abundant in certain areas, it is not exploited commercially, and, because of its relatively small size, its economic potentiality has not been investigated by U.S. Government research vessels (Harvey R. Bullis, personal communication). Recently, in the course of explorations by the Woods Hole Oceanographic Institution (Haedrich, Rowe, and Polloni, 1975), one female of this species (carapace length 17.5 mm) was taken off Barnegat, N.J., during Gosnold cruise 197, stn 111, 40°00’N, 70°47’W, depth 275-290 m. This record extends the known range of the species about 800 km north of the previously most northern precisely recorded locality, off Cape Lookout, N.C., 34°45’N, 75°28'W, depth 320 m, Combat stn 175 (Bullis and Thompson, 1965). Of greater interest has been the discovery of its presence off southern Brazil where 16 males (15 to 20 mm carapace length) and 22 females (18 to 23.5 mm carapace length) were collected by Dr. V. N. Seménov, during an exploratory cruise of the R/V Akademic Knipovitch, off Rio Grande do Sul, at 32°45'24”S, 50°24’W, depth 345-260 m (Figure 1). Hereto- fore, the southernmost locality reported for this shrimp was off Surinam, 7°38'N, 54°43'W, 457 m, Oregon stn 2008. Actually, ‘‘megalops’’ had been collected (one male, 14 females) farther southeast, north of Roche Brigan- din, French Guiana, at 7°11'N, 52°58’W, depth 457 m, Oregon stn 2028, but VOLUME 92, NUMBER 1 205 Fig. 1. Geographic distribution of Penaeopsis serrata in the western Atlantic. this find has not been previously published. The presence of P. serrata off Rio Grande do Sul represents an extension of about 6,400 km southeast of its reported southern limit (Bullis and Thompson, 1965), and an addition to the decapod crustacea of Brazil. A detailed examination of the specimens from southern Brazil revealed no significant differences between them and representatives of the species occurring north of the equator. The rostra of the Brazilian specimens are almost straight or slightly arched basally, and armed with 12 to 16 teeth in addition to the epigastric tooth, variations that fall within the range of those 206 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON a | Fig. 2. P. serrata, 2 21 mm cl, off Rio Grande do Sul, Brazil. Thelycum, ventral view. Scale = 1 mm. present in northern populations. The branchiocardiac carina and the long, interrupted cicatrix extending along the lateral surface of sixth abdominal somite are very weak in all specimens, and the petasmata are virtually identical in all males. In the Brazilian females, the thelyca are similar to those in most individuals of northern populations: the plate of sternite XIV (Fig. 2) is strongly produced anterolaterally and the median plate of XIII is subcordiform or subsemicircular. Our examination of extensive collec- tions of P. serrata from the north Atlantic has demonstrated that in the popu- lations from the Caribbean and the Atlantic coast of northern South Amer- ica, the third maxilliped and the pereopods are proportionately longer than they are in the populations from the Gulf of Mexico and the Atlantic coast of the United States. In the Brazilian specimens, these appendages extend as far as those in specimens from the Caribbean and northern South America. The VOLUME 92, NUMBER 1 207 uniformity in the Caribbean—northern South America and southern Brazil specimens is remarkable if the latter are members of a truly disjunct popu- lation. The specimens from off New Jersey and Rio Grande do Sul are deposited at the Museum of Natural History, Smithsonian Institution, catalog numbers USNM 168338 and USNM 171466, respectively. Those from New Jersey were donated by R. L. Haedrich, G. T. Rowe, and P. T. Polloni (to whom we are grateful), of the Woods Hole Oceanographic Institution, and those from off Rio Grande do Sul, by the second author. Acknowledgments We are grateful to V. N. Seménov, who collected the specimens from Brazil and made them available to us. Thanks are also extended to B. B. Collette and A. B. Williams for their critical reading of the manuscript. Illustrations are by Maria M. Diéguez. Literature Cited Bate, C. S. 1881. On the Penaeidea.—Ann. Mag. Nat. Hist., Ser. 5, 8:169-196. Bullis, H. R., Jr., and J. R. Thompson. 1965. Collections by the exploratory fishing vessels Oregon, Silver Bay, Combat, and Pelican made during 1956-1960 in the southwestern North Atlantic.—U.S. Fish Wildl. Serv., Spec. Sci. Rept. Fish. 510, 130 p. Haedrich, R. L., G. T. Rowe, and P. T. Polloni. 1975. Zonation and faunal composition of epibenthic populations on the continental slope south of New England.—Jour. Mar. Res. 33:191-212. Hay, W. P., and C. A. Shore. 1918. The decapod crustaceans of Beaufort, N.C., and the surrounding region.—Bull. U.S. Bur. Fish. 35:369-475. Maurin, C. 1961. Répartition des crevettes profondes sur les cotes sud du bassin occidental de la Méditerranée et dans la région atlantique ibéro-marocaine.—Comm. Int. Explor. Sci. Mer Médit., Rapp. P.-V. Reun. 16:529-532. . 1968. Les Crustacés capturés par la ‘‘Thalassa’’ au large des cotes nord-ouest afri- caines.—Rev. Roum. Biol., Ser. Zool. 13:479-493. Smith, S. I. 1885. On some genera and species of Penaeidae, mostly from recent dredgings of the United States Fish Commission.—Proc. U.S. Natl. Mus. 8:170-190. (IPF) National Marine Fisheries Service, Systematics Laboratory, Na- tional Museum of Natural History, Washington, D.C. 20560; (BGI) All- Union Research Institute of Marine Fisheries and Oceanography, VNIRO, Moscow, B-140, USSR. PROC. BIOL. SOC. WASH. 92(1), 1979, pp. 208-215 PENAEOPSIS JERR YI, NEW SPECIES FROM THE INDIAN OCEAN (CRUSTACEA: PENAEOIDEA) Isabel Perez Farfante Abstract.—Penaeopsis jerryi, new species, is described and illustrated. This penaeid shrimp occurs throughout the Indian Ocean, from the Bay of Bengal to Mozambique and Madagascar, at depths between 183 and 677 m. It is distinguished from its close relative P. rectacuta, occurring eastward of the Strait of Malacca, by the relative position of the antennal and hepatic spines, the length of the branchiocardiac carina, and thelycal features. In my studies of the genus Penaeopsis it became obvious that soon after Bate (1881, 1888) described the first two members of the genus from the Indo-West Pacific [P. rectacuta from the Philippines, and P. serrata (=P. challengeri de Man 1911) from the Fiji Islands] specimens of another species, occurring only in the Indian Ocean, had been repeatedly misiden- tified as P. rectacuta. The confusion in the identity of this third species has persisted for almost a century, beginning in 1891 when Wood-Mason as- signed to P. rectacuta specimens from the Andaman Sea. This formerly misidentified species, herein designated Penaeopsis jerryi, occurs in the Indian Ocean, where P. balssi Ivanov and Hassan, 1976, is also present. Three other members of the genus, P. challengeri de Man, 1911, P. eduardoi Pérez Farfante, 1977, and P. rectacuta (Bate, 1881) are found in the Indo-West Pacific, east of the Strait of Malacca (P. eduardoi was mis- takingly recorded from the Indian Ocean in the original description). The remaining species assigned to the genus, P. serrata Bate 1881, is restricted to the Atlantic. Penaeopsis jerryi is widely distributed through- out the Indian Ocean and fairly abundant in certain areas (for example, off the southwestern coast of India where exploratory fishing has in- dicated commercial potentiality), but few locality records are known and only a single one has been reported along the east coast of Africa. Fortunately, most of the references to this shrimp include descriptions of its diagnostic features and/or clear illustrations and these have been very helpful in this study. The account below is based on the examination of 85 specimens, which represent populations from three widely distant regions of the Indian Ocean: the Gulf of Aden, off the southwestern coast of India, and north of the Andaman Islands in the Bay of Bengal. The material examined is in the collections of the British Museum (Natural History), National Museum of Natural History (USNM), and Zoological Survey of India (ZSI). VOLUME 92, NUMBER I 209 Fig. 1. Penaeopsis jerryi n. sp., paratype 2 17.5 mm cl, off Berbera, Gulf of Aden, Somalia. Lateral view. Scale = 5 mm. Penaeopsis jerryi, new species Figures 1-3 Metapenaeus rectacutus. Wood-Mason 1891:274.—Alcock 1901b:50. Peneus rectacutus. Alcock 1898:73. Penaeus rectacutus. Alcock and Anderson 1899:278. Peneus (Parapeneus) rectacutus. Alcock 1901a:17.—Alcock and McArdle 1901, pl. 49, fig. 5. Parapeneus rectacutus. Alcock 1902:268, fig. 62; 1905:520 [part]; 1906:33, pl. 6, fig. 19, 19a—b.—Kemp and Sewell 1912:16. Penaeopsis rectacutus. Ramadan 1938:67, fig. 12a—b.—Sewell 1955:202.— Kurian 1964:216. Penaeopsis rectacuta. Holthuis and Rosa 1965:3 [part]—George 1966:342; 1969:27.—Longhurst 1971:224.—Starobogatov 1972:390 [key without illus- trations].—Crosnier and Jouannic 1973:12, pl. 3, fig. 3.—Ivanov and Has- san 1976:5, fig. 3. Not Penaeus rectacutus Bate 1881. Material.—Holotype: 2, 20.5 mm carapace length, 18.5 mm rostrum length, about 104 mm total length; type-locality: off Berbera, Somalia, Gulf of Aden, 10°29'48’N; 45°01'48”"E, 186 m, 21 September 1933, John Murray Expedition stn 16, BMNH 1978:325. Paratypes: 20 ¢ 49 2, BMNH 1978:326, collected with holotype; 2 6 2 2, USNM 171430, off Saihut, Yemen, 15°10’N; 50°58’E, 240-239 m, 16 May 210 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 2. Penaeopsis jerryi n.sp., holotype 2 20.5 mm cl, off Berbera, Gulf of Aden, Somalia. A, Anteroventral part of carapace; B, Telson and left uropod, dorsal view. Scales A = 2 mm, B = 5 mm. 1971, A. D. Druzhinin. 2 6 2 2, USNM 171431, off Cochin, India, summer 1978, Staff Depart. Mar. Sci. Univ. Cochin. 1 2, USNM 42755, off False Divi Point, India, 15°56’50’”N; 81°30'30’E, 439-505 m, 24 December 1890, Investigator stn 120.2 6 1 2, ZSI 2589-95/10, N of North Andaman I, 14°13'N; 93°40’E, 677-766 m, 8 April 1898, Investigator stn 235. Description.—Rostrum (Fig. 1) almost horizontal, straight or slightly sin- uous (occasionally convex basally, straight anteriorly), falling short of to overreaching distal margin of antennular peduncle, its length 0.8 to 0.9 that of carapace. Rostral plus epigastric teeth 12 to 16, second rostral tooth (sometimes apex to, at most, midlength of first) situated in line with orbital margin. Postrostral carina extending posteriorly to about level of dorsal ex- tremity of cervical sulcus; minute dorsal tubercle located near posterior mar- VOLUME 92, NUMBER 1 211 A Fig. 3. Penaeopsis jerryi n.sp., paratype 6 21 mm cl, off Cochin, India. A, Right half of petasma, dorsal view; B, Ventral view; C, Appendix masculina, dorsolateral view. Scales = 1 mm. gin of carapace. Antennal carina short but prominent. Hepatic spine slightly larger than, and situated at about same level as (rather than distinctly ventral to) that of, antennal spine; pterygostomian spine sharp. Anteroventral ex- tremity of carapace (excluding ventral membrane) forming angle of about 90° (Fig. 2A). Cervical carina sharp, accompanying sulcus well marked; hepatic carina slanting sinuously from below hepatic spine to pterygostomian spine; branchiocardiac carina strong, with anterior extremity almost reaching pos- terior end of hepatic sulcus and extending posterodorsally to near margin of carapace. Antennular flagella sexually dimorphic. In male, ventral flagellum shorter than dorsal, with proximal part forming rigid, flattened, semicircular loop bearing small basal scale and ending distally in conspicuous knob; distal part straight and somewhat compressed laterally. In female, ventral flagel- lum also bearing small basal scale, but straight along entire length and longer than dorsal. Scaphocerite falling short of to overreaching distal end of antennular pe- duncle. Antennal flagellum more than twice length of animal (Kurian 1964). 72} 92 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Abdomen with sixth somite elongate, bearing long, strong, interrupted cicatrix on lateral surface; cicatrix also on fifth and fourth somites. Telson with lateral margins usually bearing 3 pairs of small, movable spines (Fig. 2B); fixed spines variable in length, reaching at most base of distal third of terminal portion; latter narrowly hastate or with lateral margins basally rounded, and dorsal surface moderately convex. Petasma (Fig. 3A—B) with dorsomedian lobule produced into rather broad distomedian projection, and bearing elongate distal plate and broader, sub- triangular proximal plate raised mesially in low crest; rib of dorsolateral lobule terminating proximally in semicircular to subcircular process. Ven- trolateral lobule bearing distally rather flexible and translucent marginal re- gion, reflexed inwardly; distal part of ventral costa curving gradually dorsomesially and ending in short, relatively narrow process reaching ap- proximately to level of cincinnuli (description and illustrations based upon unfolded petasma). Appendix masculina (Fig. 3C) transversely oval, about 1.7 as wide as long; bearing band of setae around free margin and tuft of slightly longer setae on dorsomesial extremity. Thelycum (Fig. 4) with plate of sternite XIV subelliptical in outline, its anterior border strongly arcuate and inclined posterolaterally, and antero- lateral and posterolateral corners arched, plate sloping gently toward sub- median depressions variable in length, and bearing long marginal setae; median ridge broadest and most salient posteriorly, tapering anteriorly, sometimes reduced to posterior tubercle. Median plate of sternite XIII sub- semicircular to roughly trilobed, sometimes with minute anteromedian spine, and covered with setae except for central depression; posteromedian projection broad, with posterior margin entire or very shallowly emarginate. Sternite XII bearing posteromedian, subconical tooth with apex directed anteroventrally. Color: Red (Wood-Mason 1891), or dark brown with a reddish tint (Kurian 1964). Maximum lengths: 160 mm tl (Crosnier and Jouannic 1973). Largest spec- imens examined by me: males 23 mm cl, about 107 mm tl; females 33 mm cl, about 138 mm (cl, carapace length; tl, total length). Geographic and bathymetric ranges.—Indian Ocean, from the Bay of Bengal (Andaman Sea: Wood-Mason 1891, Alcock 1899, Alcock and An- derson 1899; off Madras: Alcock 1906), through the Arabian Sea (southwest of Cochin: Kemp and Sewell 1912) to the Gulf of Aden, (off Berbera: Ra- madan 1938), and south to off Mozambique (Ivanov and Hassan 1976) and Madagascar (Crosnier and Jouannic 1973). It has been found at depths be- tween 183 and 677 m. Affinities. —Penaeopsis jerryi differs from P. rectacuta mainly by the po- sition of the hepatic spine, the length of the branchiocardiac carina, and VOLUME 92, NUMBER I 213 i _j Fig. 4. Penaeopsis jerryi n.sp., holotype. Thelycum, ventral view. Scale = 1 mm. features of the thelycum. The petasmata of the two species are virtually identical. In P. rectacuta the hepatic spine is located at a level distinctly ventral to, instead of about the same level as that of, the antennal spine, and the branchiocardiac carina ends farther from the hepatic sulcus than it does in P. jerryi. The more dorsal position of the hepatic spine in the latter has been previously pointed out and/or depicted in illustrations presented by Alcock and McArdle (1901: pl. 49: fig. 5); Alcock (1902, fig. 62; 1906:34, pl. 6: fig. 19); Ramadan (1938:67, fig. 12a); and Ivanov and Hassan (1976:7); the longer branchiocardiac carina was pointed out by Alcock and McArdle (loc. cit.); Alcock (1906); and Ramadan (loc. cit.). None of these authors, how- ever, recognized that these two features distinguish the western from the eastern species. Regarding the thelycal features, in P. rectacuta the plate of sternite XIV is usually roughly trapezoidal, with the anterior border almost straight on each side of the posteromedian projection of sternite XIII, and the antero- lateral corners forming angles, whereas in P. jerryi this plate is roughly oval, with the anterior border arcuate and the anterolateral and postero- lateral corners arched. Furthermore, in P. rectacuta the median plate of 214 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON sternite XIII is cordiform (with acute apex) whereas in P. jerryi this plate is subsemicircular or occasionally weakly trilobed as in the specimen figured by Ramadan (1938, fig. 12b). Etymology.—This species is named for my son, Gerardo (Jerry) Canet. Acknowledgments Iam much indebted to A. A. Fincham and R. W. Ingle for loaning material and for their cooperation while working at the BMNH; to B. G. Ivanov, All Union Research Institute of Marine Fisheries and Oceanography (VNIRO) and P. Subramanian, The Marine Center of Advanced Study in Marine Biology, India, for donating specimens to the Smithsonian Institution; and to K. K. Tiwari and C. Ramakrishna (ZSI) for making available to me specimens in their institution. I further thank H. H. Hobbs, Jr., F. A. Chace, Jr., and A. B. Williams for reading and commenting on the manuscript. Maria M. Diéguez, with her usual attention to detail, prepared the drawings. Literature Cited Alcock, A. W. 1898. A summary of the deep-sea zoological work of the Royal Indian Marine Survey ship Investigater from 1884 to 1897.—Sci. Mem. Med. Off. Army India 11:1- 93. . 1901a. A descriptive catalogue of the Indian deep-sea Crustacea Decapoda Macrura and Anomala, in the Indian Museum. Being a revised account of the deep-sea species collected by the Royal Indian Marine Survey ship Investigator.—Indian Museum, Cal- cutta, 286 p. . 1901b. Zoological gleanings from the Royal Indian Marine Survey ship Investigator.— Sci. Mem. Med. Off. Army India 12:35-76. . 1902. A naturalist in Indian Seas or, four years with the Royal Indian Marine Survey ship ‘Investigator.’—John Murray, Lond. p. 1-328. . 1905. A revision of the ‘‘genus’’ Peneus with diagnoses of some new species and varieties.—Ann. Mag. Nat. Hist., Ser. 7, 16:508—532. . 1906. The prawns of the Peneus group. Catalogue of the Indian decapod Crustacea in the collection of the Indian Museum. Part III.—Macrura, Indian Museum, Calcutta, 5) p> , and A. R. S. Anderson. 1899. Natural history notes from H. M. Royal Indian Marine Survey ship ‘Investigator,, Commander T. H. Heming, R.N., commanding. Series III, No. 2. An account of the deep-sea Crustacea dredged during the surveying-season of 1897-98. —Ann. Mag. Nat. Hist., Ser. 7, 3:1-27, 278-292. , and A. F. McArdle. 1901. Illustrations of the zoology of the Royal Indian Marine Survey ship Investigator, under the command of Commander T. H. Heming, R.N.., Crustacea. Part IX, plates 49-55.—Off. Supt. Gov. Print. India, Calcutta. Bate, C. S. 1881. On the Penaeidea. Ann. Mag. Nat. Hist., Ser. 5, 8:169-196. . 1888. Report on the Crustacea Macrura collected by H.M.S. Challenger during the years 1873-76.—Rep. Sci. Results voyage H.M.S. Challenger, 1873-76, Zool. 24, 942 p. Crosnier, A., and C. Jouannic. 1973. Note d’information sur les prospections de la pente continentale malgache effectuées par le N. O. Vauban—Bathymétrie—Sédimentologie— Péche au chalut.—Doc. Sci. Centre ORSTOM Nosy-Bé, no. 42, 18 p. multigr. VOLUME 92, NUMBER 1 215 George, M. J. 1966. Ona collection of penaeid prawns from the offshore waters of the south- west coast of India——Proc. Symp. Crust. (1):337-346. . 1969. Prawn fisheries of India. Systematics-taxonomic consideration and general dis- tribution.—Cent. Mar. Fish. Res. Inst., Bull. 14:5-48. Holthuis, L. B., and H. Rosa, Jr. 1965. List of species of shrimps and prawns of economic value.—FAO (Food Agric. Organ. U.N.) Fish. Tech. Pap. 52, 21 p. Ivanov, B. G., and A. M. Hassan. 1976. Penaeid shrimps (Decapods, Penaeidae) collected off east Africa by the fishing vessel ‘‘Van Gogh’’, 2. Deep-water shrimps of the genera Penaeopsis and Parapenaeus with description of Penaeopsis balssi sp. nov.—Crusta- ceana 31:1-10. Kemp, S., and R. B. S. Sewell. 1912. The species obtained by R.I.M.S.S. ‘Investigator’ during the survey season 1910-11. Notes on Decapoda in the Indian Museum. III— Rec. Indian Mus. 7:15-32. Kurian, C. V. 1964. On the occurrence of the deep-water prawn Penaeopsis rectacutus (Spence Bate) off the Kerala coast.—Current Science (Bangalore) 33(7):216—217. Longhurst, A. R. 1971. Crustacean resources. In: J. A. Gullard, the fish resources of the ocean. FAO (Food Agricul. Organ. U.N.), pp. 206—25S. Man, J. G. de. 1911. The Decapoda of the Siboga Expedition. Part. I. Family Penaeidae.— Siboga Exped. Monogr. 39a, 131 p. Pérez Farfante, I. 1977. Penaeopsis eduardoi, a new species of shrimp (Crustacea: Penaeidae) from the Indo-West Pacific.—Proc. Biol. Soc. Wash. 90:172-182. Ramadan, M. M. 1938. «Crustacea: Penaeidae. John Murray Exped. 1933-34.—Sci. Rep. 5(3):35-76. Sewell, R. B. S. 1955. A study of the sea coast of southern Arabia.—Proc. Linn. Soc. Lond. 165:188-210. Starobogatov, Y. I. 1972. Penaeidae (Crustacea Decapoda) of the Tonking Gulf. Jn The fauna of the Tonking Gulf and conditions of life in it. Acad. Sci. U.R.R.S., Zool. Inst., Explorations of the fauna of the seas, 10(18):359-415. (In Russian). Wood-Mason, J. 1891. Phylum Appendiculata. Branch Arthropoda. Class Crustacea. In J. Wood-Mason and A. Alcock (editors). Natural history notes from H.M. Indian Marine Survey steamer ‘Investigator, Commander R. F. Hoskyn, R. N., commanding. Series IT, No. 1. On the results of deep-sea dredging during the season 1890-91.—Ann. Mag. Nat. Hist., Ser. 6, 8:269-286. Systematics Laboratory, National Marine Fisheries Service, U.S. Na- tional Museum, Washington, D.C. 20560. 2.2 ot J4e8. Teel eee asi}, THe | | Sith 9 Aiea ae ae JUV A i " hott) ah 140 U0 vei Loire jag th Cn » Nps - ete eA US eee ie iow te 4 i--at emai Oe ee i Hg we Sinead veyed Ln = $i: “a b rg os a “as wt Ve icons he ae * ae et “Pay ih awl As te s ‘eae ; 7 uM ep i) ai ane Yay eye we a) ee a bes ia it nN nt 7. INFORMATION FOR CONTRIBUTORS Content.—The Proceedings of the Biological Society of Washington contains papers bearing on systematics in the biological sciences (both botany and zoology, including paleontology), and notices of business transacted at meetings of the Society. Except at the direction of the Council, only manuscripts by Society members will be accepted. Papers will be published in English, except for Latin diagnosis/description of plant taxa which should not be duplicated by an English translation, or summary in an alternate language when appropriate. 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The establishment of new taxa must conform with the requirements of the appropriate in- ternational codes of nomenclature. Authors are expected to be familiar with these codes and to comply with them. New species-group accounts must designate a type-specimen deposited in an institutional collection. The sequence of material should be: Title, Author(s), Address(es), Abstract, Text, Literature Cited, Appendix, Tables (each table numbered with an Arabic numeral and heading provided), List of Figures (entire figure legends), Figures (each numbered and identified). Type manuscripts double-spaced throughout (including tables, legends, and footnotes) on one side of paper measuring approximately 82 x 11 inches, leaving margins of at least one inch all around. Submit a facsimile with the original and retain an author’s copy. Number pages consecutively at the top. One manuscript page = approximately % of a printed page. Underline singly scientific names of genera and lower categories; leave other indications to the editors. Figures and tables, with their legends and headings, should usually be self explanatory, not requiring reference to the text. Indicate their approximate placement by a pencil mark in the margin of the manuscript. Illustrations should be planned in proportions that will efficiently use space on the type bed of the Proceedings (11.8 x 18 cm). Legends require 4 mm of column length per line. _ Proofs.—Galley proof will be submitted to authors for correction and approval. Changes other than printer’s or editor’s errors may be charged to authors. Reprint orders will be taken with returned proof. Aa j , CS eee ee ee CONTENTS A new species of Diploglossus (Sauria: Anguidae) from Hispaniola Albert Schwartz, Eugene D. Graham, Jr., and Julian J. Duval Some phylogenetic implications of a discovery of Aspidoras pauciradiatus (Pisces: Siluriformes: Callichthyidae) from the Rio Negro in Brazil Stanley H. Weitzman and Martha H. Balph The humerus of Xenicibis, the extinct flightless ibis of Jamaica Storrs L. Olson and David W. Steadman Review of Toxotherium (Perissodactyla: Rhinocerotoidea) with new material from the early Oligocene of Wyoming Robert J. Emry A new species of Limnodrilus (Oligochaeta: Tubificidae) from Jamaica Ralph O. Brinkhurst Redescription of the thecal tabulation of Scrippsiella gregaria (Lombard and Capon) comb. nov. (Pyrrhophyta) with light and scanning electron microscopy Alfred R. Loeblich, III, James L. Sherley, and Robert J. Schmidt Poecilostome copepods (Lichomolgidae) from the alcyonacean coral Cespitularia multi- pinnata in the Moluccas Arthur G. Humes and Masahiro Dojiri Pythonaster pacificus n. sp., a new starfish of the family Myxasteridae (Echinodermata: Asteroidea) Maureen E. Downey Two new species of the genus [dunella Sars, 1895 (Crustacea: Amphipoda) with re- marks on the other species. (Contribution to the knowledge of the Amphipoda 94). Gordan S. Karaman Anatomical notes on Lutodrilus multivesiculatus (Annelida: Oligochaeta) AY Michael L. McMahan A new species of Cerapus Say, 1817 (Crustacea: Amphipoda) from the northern Gulf of Mexico, with notes on its ecology James D. Thomas and Richard W. Heard Classificatory revisions in gammaridean Amphipoda (Crustacea), Part 1. Gordan S. Karaman and J. Laurens Barnard Three new species of Indo-West Pacific lizardfish (Synodontidae) Barry C. Russell and Roger F. Cressey A reevaluation of Sesarma barbimanum Cano, 1889 and S. crassipes Cano, 1889 (Crus- tacea: Decapoda: Grapsidae) Lawrence G. Abele A new species of Spilocuma (Cumacea: Bodotriidae: Mancocuminae) from the Gulf of Mexico Paul E. Omholt and Richard W. Heard Correct citations for some North American bird taxa Richard C. Banks and M. Ralph Browning Range extension of Penaeopsis serrata (Crustacea: Penaeoidea) to off New Jersey and Rio Grande do Sul Isabel Pérez Farfante and Boris G. Ivanov Penaeopsis jerryi, new species from the Indian Ocean (Crustacea: Penaeoidea) Isabel Pérez Farfante 166 176 184 195 204 208 Proceedings of the BIOLOGICAL SOCIETY of WASHINGTON } ; j Volume 92 16 August 1979 Number 2 THE BIOLOGICAL SOCIETY OF WASHINGTON 1978-1979 Officers President: Oliver S. Flint, Jr. Secretary: Michael A. Bogan Vice President: Richard Banks Treasurer: David L. Pawson Elected Council Arthur H. Clarke W. Duane Hope Robert H. Gibbs, Jr. Catherine J. Kerby Austin B. Williams PROCEEDINGS Editor: C. W. Hart, Jr. Associate Editors Classical Languages: George C. Steyskal Invertebrates: Thomas E. Bowman Plants: David B. Lellinger Vertebrates: Richard Banks Insects: Robert D. Gordon Membership in the Society is open to anyone who wishes to join. There are no prerequisites. Annual dues of $7.00 include subscription to the Proceedings of the Biological Society of Washington. Correspondence concerning membership should be addressed to the Treasurer, Biological Society of Washington, National Museum of Natural History, Smithsonian Insti- tution, Washington, D.C. 20560. The Proceedings of the Biological Society of Washington is issued quarterly in February, May, August and November. Manuscripts, corrected proofs, editorial questions should be sent to the Editor, Biological Society of Washington, National Museum of Natural History, Smithsonian Institution, Wash- ington, D.C. 20560. Known office of publication: National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560. Printed for the Society by Allen Press, Inc., Lawrence, Kansas 66044 Second class postage paid at Washington, D.C., and additional mailing office. PROC. BIOL. SOC. WASH. 92(2), 1979, pp. 217-224 NEW RECORDS OF THE SPECIES OF GLYPHOCRANGON IN THE NORTHEASTERN PACIFIC OCEAN (CARIDEA: GLYPHOCRANGONIDAE) Mary K. Wicksten Abstract.—Four species of Glyphocrangon are recorded from the north- eastern Pacific. Glyphocrangon spinulosa and G. vicaria are reported for the first time from southern California. Glyphocrangon sicaria has been found off Punta Guiones, Costa Rica. Illustrations and a key are provided for the species. Members of the genus Glyphocrangon are benthic bathyal shrimps. AI- though the four species in the northeastern Pacific Ocean were described by Faxon (1893, 1896) and included in a review of the family Glyphocran- gonidae by De Man (1921), none have been reported previously from Cali- fornia or the outer coast of Baja California, Mexico. Glyphocrangon vicaria has not been illustrated, although Faxon (op. cit.) described it in detail. Recent collections made by the R.V. Agassiz, Melville, Oconostota, and Spencer F. Baird of Scripps Institution of Oceanography and the R.V. Ve- lero IV of the Allan Hancock Foundation included G. sicaria, G. spinulosa, and G. vicaria. Material of G. alata taken by the R.V. SNP-/ was donated to the Allan Hancock Foundation. Examination of these specimens has revealed extensions of the ranges and has allowed comparison of G. vicaria with other species. A key to the species of Glyphocrangon in the northeastern Pacific Ocean is provided. I follow the names proposed by Holthuis (1971) for the spines and carinae. Measurements are in mm. I thank Fenner A. Chace, Jr., National Museum of Natural History, for the loan of specimens of G. vicaria, G. nobilis, and G. longirostris; and Spencer R. Luke, Scripps Institution of Oceanography for the loan of spec- imens of G. sicaria, G. spinulosa, and G. vicaria. The specimens taken by the R.V. Velero IV at station 25654 were collected during cruises supported by the U.S. Office of Naval Research, contract number N00015-75-C-0683, Richard E. Pieper, chief investigator. The specimens taken at stations SNP1-13, 24, 27, and 28 were collected during the Peruvian Coastal Islands Investigations, sponsored by the Los Angeles County Museum of Natural History. Teri A. Hoffman prepared the drawings. This paper is contribution number XXX of the Allan Hancock Foundation. 218 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Glyphocrangon alata Faxon 1893 Fig. 1A, B, C Glyphocrangon alata Faxon, 1893:201; 137-138, pl. 37.—De Man, 1921:214.—Bahamonde, 1963:4.—del Solar, 1972:10. Previous records.—Off Acapulco, Mexico (16°33’N, 99°53’W), 219 m, brown sand, 11 March 1891, Albatross sta. 3418. Gulf of Panama (7°31'N, 78°39'W), 1350 m, rocky bottom, 11 April 1891, Albatross sta. 3395 (Faxon, 1895). Vicinity of Valparaiso, Chile, 600 m (Bahamonde, 1963). Off Ecuador (3°47'S, 81°12’ W), 650 m, 17 March 1971, R.V. Wiracocha. 7°26'S, 80°46’ W, 839 m; 10°51'S, 78°30’ W, 680 m; 18°19’S, 71°12'W, 810 m; 1970-1972, ship SNP-I (del Solar, 1972). Material examined.—PERU: Between Lobos de Tierra and Lobos de Afuera (6°42'S, 80°59’W), 800 m, 22 January 1974, SNP1-24, 2 specimens. Near Lobos de Afuera (7°7'S, 80°46’W), 1200 m, 20 January 1974, SNPI- 13, 1 specimen. 55 mi. S of Lobos de Afuera (7°44’S, 80°30’ W), 750-760 m, 23 January 1974, SNP1-28, 11 specimens, 4 ovigerous. 50 mi. SE of Lobos de Afuera (7°46’S, 80°31’W), 800 m, beam trawl, 23 January 1974, SNPI- 27, 11 specimens, 1 ovigerous. Size distribution.—Largest individual: Total length (tip of rostrum to tip of telson), 102; rostrum, 17; carapace, 31; abdomen without telson, 50, telson broken. Smallest individual: total length, 39; rostrum, 7; carapace, 10; abdomen, 15; telson, 7. Number of eggs per female.—28-S1. Remarks.—No other specimens than those reported by Faxon (1895) have been reported from the northeastern Pacific Ocean. Glyphocrangon sicaria Faxon 1893 Fig. 2A, B Glyphocrangon sicaria Faxon, 1893:202—203;—1895:144—-146, pl. 39.—De Man, 1921:215. Previous record.—Gulf of Panama (6°21'N, 80°41’W), 3310 m, green mud, 7 March 1891, Albatross sta. 3382 (Faxon, 1895). Material examined.—South of Punta Guiones, Costa Rica (9°35’N, 85°41'W), 1454 m, 10-foot Isaacs-Kidd midwater trawl, 20 April 1973, R.V. Agassiz, 1 specimen. Size.—Total length, 31; rostrum, 8; carapace, 8; abdomen, 10; telson, 5. = Fig. 1. Glyphocrangon alata. Female, total length 88 mm. South of Lobos de Afuera, Peru : A, Dorsal view; B, Lateral view of front; C, Dorsal view of front. S indicates prominent spine. VOLUME 92, NUMBER 2 220 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON = Coes ay) eas , ee )) / (a aS IN ES en S\y WY) aw) ST Q Ni B y, ) NS : OSmm , Fig. 2. Glyphocrangon sicaria. Juvenile, total length 31 mm. South of Punta Guiones, Costa Rica. A, Dorsal view; B, Lateral view. Glyphocrangon spinulosa Faxon 1893 Fig. 3A, B Glyphocrangon spinulosa Faxon, 1893:202; 1895:138—140, pl. 38.—De Man, 1921:215. Previous records. —Gulf of California (26°48’N, 110°45’W), 1586 m, brown mud, 22 April 1891, Albatross sta. 3435. Near Las Tres Marias Islands (21°15'N, 106°23'’W), 1248 m, gray sand, 18 April 1891, Albatross sta. 3424. Off Acapulco (16°34’N, 100°3’W), 1374 m, green mud, 11 April 1891, A/- batross sta. 3419. Off Acapulco (16°33’N, 99°53’W), 1218 m, sand, 11 April 1891, Albatross sta. 3418. Off Mariato Point (7°7'N, 80°34’ W), 1283 m, green mud, 23 February 1891, Albatross sta. 3353 (Faxon, 1895). Material examined.—Cortez Basin (32°32’N, 118°53’W to 32°33’N, 118°54'W), 1298 m, 25-foot otter trawl, 6 May 1975, R.V. Agassiz sta. B175- 2, 1 specimen. Cabo San Lucas Canyon, Baja California, Mexico; 1097 m, rock dredge, January 1957, Spencer F. Baird cruise dredge sta. 2, 1 speci- men. Also 12 specimens from 5 other stations from the Gulf of California, Mexico; to Costa Rica, 1157-1875 m, 1967-1973. Size distribution.—Largest individual: Total length, 110; rostrum, 17; car- apace, 31; abdomen, 43; telson, 17. Smallest individual: total length, 53; rostrum, 11; carapace, 13; abdomen, 19; telson, 10. Number of eggs per female.—11-16. VOLUME 92, NUMBER 2 Fig. 3. Glyphocrangon spinulosa. Female, total length 95 mm. 14 miles from Punta Guiones, Costa Rica. A, Dorsal view; B, Lateral view. Glyphocrangon vicaria Faxon 1896 Fig. 4A, B Glyphocrangon nobilis A. Milne Edwards?: Faxon, 1895:142-144. Glyphocrangon vicaria Faxon, 1896:159.—De Man, 1921:215. 222 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 4. Glyphocrangon vicaria. Female, total length 160 mm. Off Baja California, Mexico. A, Dorsal view; B, Lateral view. Previous records.—11 stations, from the Gulf of California (23°59’N, 108°40’W) to between Galera Point and the Galapagos Islands (0°36’N, 86°47'W), 1374-2441 m, February—April 1891, R.V. Albatross (Faxon, 1895). Material examined.—SAN CLEMENTE BASIN, CALIFORNIA: 32°35'’N, 118°14’W, 1628-1838 m, 25-foot otter trawl, 12 August 1971, R.V. Agassiz sta. B171-34, 10 specimens, 6 ovigerous. 32°28'’N, 118°7’W to 32°29'N, 118°6'W, 1829 m, 25-foot otter trawl, 5 May 1975, R.V. Agassiz sta. B175-1, 12 specimens, 3 ovigerous. 32°29’N, 118°8’W, 1792-1866 m, 40- foot otter trawl, 16 September 1971, R.V. Agassiz sta. B171-37, 45 speci- mens, 8 ovigerous and 2 with parasitic isopods (Munidion sp.). 32°28'N, 117°52’'W, 1527-1545 m, 16-foot otter trawl, 5 September 1974, R.V. Ocono- stota sta. B174-30, 1 specimen. 32°27’N, 117°56’W to 32°34'N, 118°8’W, 938 m, Isaacs-Kidd midwater trawl, 26 January 1977, R.V. Velero IV sta. 25654, 5 specimens. 32°26’N, 117°49'W to 32°29’N, 117°58'W, 1750 m, 25- VOLUME 92, NUMBER 2 223 foot otter trawl, 27 July 1973, R.V. Agassiz sta. B173-63, 1 ovigerous fe- male. SAN DIEGO TROUGH, CALIFORNIA: 32°33'N, 118°6’W, 1829 m, 40- foot otter trawl, 3 November 1971, R.V. Agassiz sta. B171-40, 1 specimen. 32°29'N, 117°31'W, 1225-1244 m, 40-foot otter trawl, 14 September 1971, R.V. Agassiz sta. B171-36, 1 specimen. BAJA CALIFORNIA, MEXICO: Abyssal plain (31°24’N, 120°12’W), 3880 m, 25-foot otter trawl, 21 March 1970, R.V. Melville sta. B170-38, 1 ovigerous female. In addition, the following specimens were examined: 50 from 10 additional stations taken from off the west coast of Baja California, Mexico; 3 taken by the R.V. Albatross at station 3413 (between the Galapagos Islands and Acapulco, 2°34’N, 92°6’W); and 2 from off Punta Guiones, Costa Rica (R.V. Velero IV sta. 18932). Size distribution.—Largest individual: Total length, 157; rostrum, 26; car- apace, 43; abdomen, 65; telson, 23. Smallest individual: total length, 28; rostrum, 6; carapace, 7; abdomen, 10; telson, 5. Number of eggs per female.—10-29. Remarks.—In his comments on this species, Faxon (1895) mentioned that the animal closely resembled the Atlantic species G. nobilis A. Milne-Ed- wards 1881 and G. longirostris (Smith 1882). However, G. vicaria has a notch posterior to the anterior spine of the anterior sublateral carina, which the two Atlantic species lack. The spines and other sculpturing of the car- apace and abdomen of G. vicaria are much more prominent than in G. nobilis. The corrugation of the rostrum of G. vicaria is weaker than in G. longirostris. As seen from the dorsal aspect, the branchiostegal spine of G. vicaria is exposed rather than partly obscured by the antennal spine as in G. nobilis and G. longirostris. The specimens of G. vicaria that were examined agree well with the description given by Faxon (1895, 1896) and the specimens from Albatross station 3413. The corrugation of the rostrum is difficult to see, being rep- resented in some individuals by a series of feathery lines along the dorsal midrib. The rostrum is more lanceolate in smaller individuals than in large ones. The hepatic tubercle on the antennal carina of the carapace and the tubercles of the submedian and intermediate carinae may be eroded in older animals instead of being sharply spiniform, but the sculpturing is observable readily on careful examination. Taken with G. vicaria at Velero IV stations 13770 and 18932 were benthic decapods including galatheid anomurans (Munidopsis diomedeae [Faxon 1895] and Munidopsis verrilli Benedict 1902), hermit crabs (Parapagurus pilosimanus benedicti de Saint Laurent 1972), pandalid shrimp (Heterocar- pus hostilis Faxon 1893) and broken back shrimp (Lebbeus washingtonianus 224 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON [Rathbun 1902]). Like G. vicaria, these animals probably inhabit soft sub- strates. KEY TO THE NORTHEASTERN PACIFIC SPECIES OF GLYPHOCRANGON 1. Anterior lateral carina of carapace ending in prominent anterior spine between antennal and branchiostegal spines........... G. alata — Anterior lateral carina of carapace not ending in prominent anterior spine between antennal and branchiostegal spines................ y, 2. Rostrum with 6-7 teeth, carapace covered with dense spinules . Aas al ae sus ee har flare oda Rie desicay nh Malar oir ,-t otemihe- ei ambyey ‘ac G. jniMona — Rostrum with 2 pairs of spines, carapace vith spines or tubercles instead’ of ‘Spimules...'.. eo PR es See ee ae 3 3. Two hepatic tubercles. Rostrum with 2 pairs of spines, second of them obsolete, without corrugation....................05. G. sicaria — One hepatic tubercle. Rostrum with 2 well-developed spines, with COMUGation’ 25 PELL Ae PII Nan AED Dae ie G. vicaria Literature Cited Bahamonde, N. 1963. Decapodos de la fauna preabismal de Chile.—Noticiaro mensual del Museo Nacional de Historia Natural, Santiago (Chile) 7(81):1—10. 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 . . . by the U.S. Fish Commission steamer ‘‘Albatross,’’ during 1891 .. . VI. Preliminary descrip- tions of new species of Crustacea.—Bull. Mus. Comp. Zool., Harvard Univ. 24(7):149- 220. Faxon, W. 1895. Reports on an exploration off the west coast of Mexico, Central and South America, and off the Galapagos Islands ... by the U.S. Fish Commission steamer ‘‘Albatross,’’ during 1891 ... XV. The stalk-eyed Crustacea—Mem. Mus. Comp. Zool., Harvard Univ. 18:1—292. Faxon, W. 1896. Reports on the results of dredging under the supervision of Alexander Agassiz, in the Gulf of Mexico and the Caribbean Sea, and on the east coast of the United States, 1877-1880, by the U.S. Coast Survey steamer ‘‘Blake’’ . . . 37. Supple- mentary notes on the Crustacea.—Bull. Mus. Comp. Zool., Harvard Univ. 30(3):153- 166. Holthuis, L. B. 1971. The Atlantic shrimps of the deep-sea genus Glyphocrangon A. Milne- Edwards, 1881.—Bull. Mar. Sci. 21(1):267-373. Man, J. G. De. 1921. The Decapoda of the Siboga Expedition. Part IV. Families Pasiphaeidae, Stylodactylidae, Hoplophoridae, Nematocarcinidae, Thalassocaridae, Pandalidae, Psal- idopodidae, Gnathophyllidae, Processidae, Glyphocrangonidae, and Crangonidae.—Si- boga Expeditie 34a*. 318 pp. Solar, E. del. 1972. Addenda al catalogo de Crustaceos del Peru.—Inst. del Mar del Peru Informe no. 38. 21 pp. Allan Hancock Foundation, University of Southern California, University Park, Los Angeles, California 90007. PROC. BIOL. SOC. WASH. 92(2), 1979, pp. 225-252 THE CRAYFISHES OF NEW ENGLAND Denton W. Crocker Abstract.—Ten crayfish species have been collected in New England. On the basis of the known limits of their geographic distribution elsewhere, 3 species, Procambarus (Scapulicambarus) clarkii (Girard), Orconectes rus- ticus (Girard), and O. obscurus (Hagen) have been introduced into the area by man. The time, place, and persons involved in the introductions are unknown. For 3 other species, O. limosus (Rafinesque), O. immunis (Ha- gen) and O. virilis (Hagen), though a natural entry into New England can be postulated (with greatest confidence for O. limosus), there probably has been considerable transfer inter- and intraregionally by man. Four species have distributions in New England which largely have been attained natu- rally: Cambarus (Cambarus) bartonii (Fabricius), C. (Puncticambarus) ro- bustus Girard, O. propinquus (Girard), and Procambarus (Ortmannicus) acutus acutus (Girard). Life history information is tabulated. A systematic list, figures, and distribution maps for each species are provided. The presence of crayfishes in New England, with species unidentified, has been recorded by historians and essayists (Williamson, 1832:165; Tho- reau, 1864:237). Information on the distributions of particular species began to accumulate with the recording of Astacus bartonii (now Cambarus bar- tonii) in Massachusetts by Gould (1841:330) and in Vermont by Thompson (1842:170). Hagen’s monograph of 1870 adds to the distributional picture of C. bartonii in New England by including the Lake Champlain drainages in Vermont. He lists Maine, New Hampshire, Connecticut, and Rhode Island as being without crayfishes. Walter Faxon’s several major crayfish studies and lists, while taxonomically important, add relatively little to the distri- butional picture in New England. He confirms the earlier records of C. bartonii in Massachusetts and Vermont, and adds Maine (1885a:143; 1885b:158—159). In his last work on crayfishes (1914) he adds Cambarus affinis (now Orconectes limosus) to the crayfish fauna of Massachusetts (Essex and Berkshire counties, pp. 372—373) and Cambarus immunis (now Orconectes immunis) to Massachusetts and New Hampshire (p. 378). He is puzzled by the records of O. limosus in Massachusetts but gives anecdotal evidence for its introduction by man into Berkshire Co. After several pages of discussion, he concludes that the presence of O. immunis in Berkshire Co., Massachusetts, is natural, but that the New Hampshire and other Mas- sachusetts records represent introductions by man. 226 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Scattered reports over the following 45 years have added species localities and a few new state records, but the distribution in New England of all these species has remained poorly delineated. Findings of these reports are summarized briefly as follows: Rathbun (1905), C. bartonii in Connecticut (?), Maine, Massachusetts, Rhode Island (?), and Vermont; Norton (1909), C. bartonii in Maine; Osburn (1912), O. limosus introduced into Connecticut; Creaser (1933), O. immunis in Mas- sachusetts and New Hampshire; Webster (1944), O. immunis and O. Ii- mosus in Connecticut; Crocker (1957), O. limosus in Maine; Crocker and Barr (1968:73), O. propinquus in Vermont; Camougis and Hichar (1959), O. virilis in Massachusetts. Two more recent and more extensive studies are those of Aiken (1965) who studied O. immunis, O. limosus, and O. virilis in lakes and ponds in New Hampshire, and of Bell (1971) who gives new state records for C. robustus in Connecticut and for O. obscurus and O. virilis in Vermont. Hobbs (1974b:53) gives the range of Procambarus a. acutus as (in part), ‘‘Coastal plain and piedmont from Maine to Georgia... .’’ Thus Maine is clearly cited (although without a specific locality) and other New England states by inference are possible components of its range. He also ) (p. 43) includes Maine in the natural range of O. virilis. These cited works place 7 crayfish species in the 6 New England states as follows: CONNECTICUT—C. bartonii (2), C. robustus, O. immunis, O. limosus MAINE—C. bartonii, O. limosus, O. virilis, P. a. acutus MASSACHUSETTS—C. bartonii, O. immunis, O. limosus, O. virilis NEw HAmMpPSHIRE—O. immunis, O. limosus, O. virilis RHODE ISLAND—C. bartonii (?) VERMONT—C. bartonii, O. limosus, O. obscurus, O. propinquus, O. vi- rilis This is the framework on which I have attempted to round out the species patterns of distribution. I have collected or have been given a total of 10 species of crayfishes from 274 collecting sites in New England. All speci- mens on which these records are based are in the collections of the National Museum of Natural History (NMNH). Another 63 species localities have been taken from the literature. These 337 localities are represented in Fig. 2. The coverage of the 6 states and of areas within them is uneven by reason of collecting difficulties, source of gift specimens, location of my base of operations, and some conjecture as to where the areas of interest might be. Collecting has been done largely by hand picking but with the use of a 10 ft seine where conditions permit and when an assistant is available. VOLUME 92, NUMBER 2 227 Systematic List of the Crayfishes of New England Hobbs (1974a:1—4) places the families of crayfishes in the astacuran In- fraorder Astacidea, Superfamily Astacoidea. See also his checklist of North and Middle American crayfishes (Hobbs, 1974b) for full synonymies. Family Cambaridae Hobbs, 1942 Subfamily Cambarinae Hobbs, 1942 Genus Cambarus Erichson, 1846 Subgenus Cambarus Erichson, 1846 Cambarus (Cambarus) bartonii (Fabricius), 1798. [C. (C.) bar- tonii is considered now to be without subspecies following Bouchard (1975:587).] Subgenus Puncticambarus Hobbs, 1969 Cambarus (Puncticambarus) robustus Girard, 1952 Genus Orconectes Cope, 1872 Orconectes immunis (Hagen), 1870 Orconectes limosus (Rafinesque), 1817 Orconectes obscurus (Hagen), 1870 Orconectes propinquus (Girard), 1852 Orconectes rusticus (Girard), 1852 Orconectes virilis (Hagen), 1870 Genus Procambarus Ortmann, 1905. (Proposed as subgenus by Ort- mann, raised to generic rank by Hobbs, 1942.) Subgenus Ortmannicus Fowler, 1912 Procambarus (Ortmannicus) acutus acutus (Girard), 1852 Subgenus Scapulicambarus Hobbs, 1972 Procambarus (Scapulicambarus) clarkii (Girard), 1852 Identification of species can be accomplished relatively easily by com- paring specimens with the drawings (Fig. 1), supplied by H. H. Hobbs, Jr. Sexually mature male crayfishes undergo a seasonally cyclical change in the Shape of the copulatory appendages (first pleopods). Form I males are ca- pable of mating successfully. Form II males cannot do so and their first pleopods more nearly resemble those of immature males. Both forms are illustrated in Fig. 1. The copulatory appendages together with the female’s Es Fig. 1 (pp. 228 and 229). The 10 species of crayfishes in New England. With the exception of Procambarus (Ortmannicus) acutus acutus and P. (Scapulicambarus) clarkii the sequence of figures from left to right is the same: 1, Dorsal view of carapace of male, form I; 2, First pleopod of male, form I, mesial view; 3, same, lateral view; 4, same, form II, lateral view; 5, Annulus ventralis (seminal receptacle) of female; 6, Dorsal view of right chela of male. For the 2 Procambarus species, the third drawing is 2 enlarged views of the first pleopod of a first form male, lateral view above, mesial view below. 228 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Vaan aN Orconectes immunis my CG Orconectes obscurus VOLUME 92, NUMBER 2 229 GaN Procambarus (Ortmannicus) acutus acutus Cambarus (Puncticambarus) robustus SNS, Orconectes limosus 4 ZFS 230 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Table 1. Seasonal events for males of Orconectes propinquus in New England. June July August quarter 2 3 1 y 3 4 form I 0 0 Z 0 30 4 form II 1 16 0 0 0 0 annulus ventralis, into which the sperm mass is introduced, are especially useful in distinguishing among species. Life History Information Although the emphasis of the study is distribution, collections of cray- fishes contain life history information which, when added to other infor- mation, contributes to a knowledge of their total biology. Some life history information and an entry into the literature for several of the New England species occurring in New York and Ontario is given by Crocker (1957) and Crocker and Barr (1968). A current summary of the population dynamics of crayfishes and their role in ecosystems is given by Momot ef al. (1978). The following data are summarized from all collections taken in New England over a period of 20 years. Thus, for several species, the data rep- resent an average of the effects of climatic factors over the entire region and of variations in seasonal weather during the years when collecting was done. Thus the data may not necessarily represent precisely the timing of life cycle events in a particular locality in a given year. A population of a species in southern New England or a warmer body of water would be expected, for example, to lay eggs earlier than populations of the same species in more northern parts or colder water. Also, the time of seasonal molt would be less extended in any one particular locality or any one year than over the region as a whole, or over the entire period of collecting. Too few adult males of the following species were captured to permit gaining information about seasonal molting: Cambarus bartonii, C. robus- tus, Orconectes immunis, O. rusticus, Procambarus a. acutus, and P. clar- kit. Females of these species with eggs or young were taken as follows: Table 2. Seasonal events for males of Orconectes limosus in New England. June July August quarter 1 2 3 4 1 2 3 4 1 ?2 3 4 form I 2 2 1 1 0 0 0 2 4 2 eal VOLUME 92, NUMBER 2 231 Table 3. Seasonal events for males of Orconectes virilis in New England. June July August quarter 3 4 1 2 3 4 1 2 3 4 form I 3 2 1 0 2 5 3 1 9 29 form II 3 5 4 1 9 10 7 5 2 9 Orconectes immunis: 2 females with eggs, Rhode Island, 2 May 1966 and 31 May 1958. Aiken (1965:242) reports that all female O. immunis col- lected from Newfound Lake, New Hampshire, on 13 and 14 June 1963 were carrying young. O. rusticus: 2 females with young, Maine, 11 June 1959. Procambarus a. acutus: 1 female with young, Rhode Island, 12 Aug 1971. An extensive bibliography for this species and for P. clarkii is given by Spohrer et al. (1975). For the remaining 3 species the data, though far from satisfactory, are more suggestive of seasonal events. Orconectes propinquus: Seasonal events for males are given in Table 1. The earliest spring form II] male was taken 11 June 1953 in Massachusetts. The taking of 30 males in the third week of August, all of them form I, suggests that the majority of individuals are form I by that time. I have no records of females with eggs or young from New England. Orconectes limosus: Seasonal events for males are given in Table 2. The lack of collecting in March, April, and May prevents documenting the time of the spring molt, but the data strongly suggest that the late summer molt occurs among most individuals during the last 2 weeks of August. Individual adult males taken in months other than those tabulated above are all form I (April—1, September—9, October—11). A female with eggs was collected in Rhode Island on 9 May 1965. Aiken (1965:241) reports taking several egg- bearing females from the Newfound River, N.H. on 13 June 1963. Orconectes virilis: Seasonal events for males are given in Table 3. Again, data are suggestive only for the late summer molt of adult males. As with O. limosus, it appears that this molt occurs most frequently in the last two Table 4. Seasonal events for males of Orconectes virilis in New Hampshire (data from, Aiken, 1965:242-243). 24 July— i9june 22 June’ ..27 June 7 July 20 Aug 27 Aug 2 Sept % form I 93 91 55 39 28 85 93 % form II A 9 45 61 72. 15 7 232 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON weeks of August. Individual adult males taken in months other than those tabulated are all form I (April—2, September—2, October—12). Aiken (1965:242—243) reports for this species the results of trapping in Lake Winnipesaukee, N.H. during the spring, summer and fall of 1962. He gives the actual number trapped only for the 24 July—20 August sample (938 males). For this locality the data summarized in Table 4 show that the early summer molt occurs primarily in the last week of June and that the late summer molt occurs in the last week of August. I have four dates for the capture of females with eggs or young as follows: female with eggs, 24 April 1952 (N.H.), 2 May 1966, and 12 May 1965 (R.1I.); 4 females with young, 25 June 1959 (Maine). Introductions and Transfers Three species occur in New England as isolated populations so far re- moved from the known geographic limits of the main distributional area of their conspecifics that I can only attribute their presence in New England to introduction by man. Orconectes obscurus: Insofar as one may judge from published data, O. obscurus has as its natural range eastern Ohio, western Pennsylvania, West Virginia, and the Allegheny, Genesee, and upper Mohawk rivers in New York (Crocker and Barr, 1968:84). The New England localities now reported for the first time (Fig. 3) are (1) Greenwater Pond and Goose Pond, each only a few miles from East Lee, Berkshire Co., Massachusetts, and (2) Lake Webb in Franklin Co., Maine. I made the Massachusetts collections on 25 August 1952 after finding in 1950 or 1951, 5 dry apparently uncataloged specimens of this species in the Museum of Comparative Zoology, Cam- bridge, Massachusetts. These are 3 males I, 1 male II, and one female. The label reads *‘Goose Pond, East Lee, Mass./Rev. Robt. Keating Smith/July, 1917.’’ The specimens were obtained from the Maine locality by me on 22 August 1956 and by Douglas Mathieu, a collecting assistant, on 6 August 1959. The region between the localities in the upper Mohawk in New York and in East Lee, Massachusetts, has been well sampled for crayfishes (see Crocker, 1957, figures 4 and 5 on pages 73 and 78), but O. obscurus has not been collected there. Bell (1971:16) reported taking O. obscurus at Hartland, Vermont, on the Connecticut River. Crocker and Barr (1968:81—84) recorded the introduction of O. obscurus into southeastern Ontario. Orconectes rusticus: This species does not quite reach Pennsylvania at the easternmost limit of its natural distribution. I cannot explain the follow- ing New England localities (Fig. 4) except as introductions by man—4 lo- VOLUME 92, NUMBER 2 233 LEGEND BOUNDARIES NATIONAL & STATE —--——-—~ COUNTY 9 —-—-—~ WATERSHEDS = -------- SCALE OF MILES ° 5 20 30 40 so (see note) a 5 S SOURCE- Traced (except drainage divides) from a ( ed 9 “THE NATIONAL SURVEY MAP OF NEW ENGLAND" 5 The National Survey, Chester, Vt. 05143 ; : : pe FX \ ry al To sr Fig. 2. Collection sites for crayfishes in New England. Dark circles = collections in Nation- al Museum of Natural History; half dark circles = literature reports of species localities; open circles = locations where at least 0.5 hr of collecting yielded no crayfishes. 234 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON calities of which occur in the Connecticut River system, and all in or within 5 mi of the mainstream of the river itself: (1) Conn. R. at N Walpole, Cheshire Co., N.H. (2) Conn. R. at N Thetford, Orange Co., Vt. (3) Wells R. about 1.5 mi NW of town of Wells River, Orange Co., Vt. (4) Stony Brook, S Hadley, Hampshire Co., Mass. (This locality was discovered independently by Douglas G. Smith of the Museum of Zoology, Univ. Mass., in collecting between 1972 and the present.) (5) Fenton River, Mansfield, Tolland Co., Conn., Thames River system. (6) Great Pond, North Pond, and Long Pond, Kennebec River drainage in Kennebec-Somerset Co., Maine. (7) Flying Pond in town of Vienna, Kennebec Co., Maine, Androscoggin River drainage. The Massachusetts, Connecticut, and Maine collections were made be- tween 1952 and 1959; those from New Hampshire and Vermont were made in 1970. I report here also for the first time the collection in 1968 of O. rusticus in New York: artificial pond about 1.25 mi N of Rynex Corners (about 6 mi W of Schenectady), Schenectady Co., Mohawk-Hudson River drainage. Crocker and Barr (1968:88—90) record the introduction of O. rusticus into southeastern (4 localities) and western (1 locality) Ontario. Procambarus clarkii: A single juvenile female of this species has paen identified by H. H. Hobbs, Jr. in a collection from University of Rhode Island: Outlet of Arcadia Pond, Mike MacCrae, 5 April 1970. This site is about 0.5 mi N of Arcadia, Washington Co., R.I., at an edge of the Arcadia Management Area and near the Arcadia Warm Water Research Station. Two other URI collections from this area contain P. a. acutus. The range of P. clarkii is given by Hobbs (1974b:65) as northern Mexico to Escambia Co., Florida, and N to southern Illinois. Transfers.—Through conversations in the summer of 1969 with Dr. Don- ald Mairs of the Maine Department of Inland Fisheries and Game, and with Mr. Robert Knowlton of the New Hampshire Fish and Game Department, I have learned that transfers of crayfishes between lakes within each of these states have been undertaken. I have been unable to obtain data on numbers and species transferred in New Hampshire, but in Maine, both O. virilis and O. limosus have been transferred between lakes in at least 6 instances, the numbers ranging from 36 to 4300. It is most likely that O. virilis is the species transferred in New Hampshire. Therefore, seemingly aberrant distributions of O. virilis in both states and O. limosus also in Maine may be the result of these attempts to establish crayfishes where they formerly were rare or absent. VOLUME 92, NUMBER 2 235 Interpretation of Distributional Data The locality records for Orconectes rusticus, O. obscurus, and Procam- barus clarkii have been explained as introductions by man. It is difficult to assess how much influence introductions have had on the current distribu- tions of other species. Though some recent introductions and transfers have been documented and though further interviewing and correspondence may gain more information, there is small likelihood that records survive, if they were kept at all, of earlier attempts and their success. The following quo- tations offer tantalizing suggestions of former crayfish abundance, a decline, and a later increase due to introduction. Documentation is not given for the general statements, however. .. . why is it that in New England crayfish are almost altogether absent .. . [Andrews, 1906a:100]. A crayfish from the Charles River, collected and contributed by Mr. Alden Cheever adds to the evidence that this animal is increasing in Massachu- setts waters. Crayfishes were once common in eastern New England, but as natives they almost disappeared. Their presence today is largely due to their introduction as food for fishes [Sanford, 1932:18]. For many years before the government stocked the ponds with crayfishes, as food for bass and other game fishes, these crustaceans were not com- mon in New England, especially east of the Connecticut River. They are now breeding rapidly, however, and it is interesting to note that their mud houses, erected on the banks of ponds and streams, but connected with the water by underground passages, are becoming more numerous. Lewis Babbit has contributed some of these crayfishes, collected in Connecticut [Sanford, 1936:11]. ... Prof. E. P. Larkin informs me that about forty years ago crayfishes (C. Bartonii?) were not uncommon at Westerly, R.I. ... [Faxon, 1885b:62]. Faxon (1885b:98) reports that O. virilis and O. immunis are two of the crayfish species most esteemed as food and that they are sometimes sent to the New York market from Milwaukee and other western cities. He also says (p. 89), ‘‘C. affinis [=O. limosus] is the common crayfish exposed to sale in the markets of New York and other eastern cities.’’ Andrews (1906b) gives the sources of crayfishes shipped to New York City (and I would conjecture to New England cities as well) as being Potomac River, Mon- treal, Milwaukee, and possibly Chicago. It seems a logical assumption that industrialization in the latter half of the nineteenth century reduced crayfish populations in parts of New England and that sport fishing, gastronomic and 236 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON ae ae 7 LEGEND BOUNDARIES NATIONAL & STATE ---»—=-=-— COUNTY WATERSHEDS SCALE OF MILES SOURCE: Traced (except drainage divides) from 4 “THE NATIONAL SURVEY MAP OF NEW ENGLAND" The National Survey, Chester, Vt. O5143 6rs0 or Fig. 3. Species localities for 4 species of crayfishes in New England. Black circles = Cambarus bartonii; circles half dark at left = C. robustus; circles half dark below = Procam- barus a. acutus; circles half dark at right = Orconectes obscurus. VOLUME 92, NUMBER 2 ACY other interests would work to replace them. Unfortunately, the total number of collections of New England crayfishes prior to 1952 either reported in the literature or on deposit in MCZ or NMNH is too small to permit docu- menting this assumption. Although the preceding information demands caution in the interpretation of distributions, I shall assume, unless specific known instances of intro- duction suggest the contrary, that distributions are natural if they have an inherent consistency and/or if Pleistocene glacial events and other geological data offer reasonable explanations for them. Procambarus (Ortmannicus) acutus acutus (Girard) Populations of crayfishes now referred to this subspecies and occurring naturally from Massachusetts to Georgia, formerly were referred to Pro- cambarus blandingii blandingii (Harlan). Hobbs (1974b:53—54 and personal communication) now restricts P. blandingii to South Carolina and southern North Carolina, and he locates P. a. acutus as now understood, not only in the coastal plain and piedmont area referred to above, but also from the Florida panhandle westward to Texas and northward to the SE corner of Minnesota. It thus has an extensive range on both sides of the Appalachian chain. New England localities are shown in Fig. 3. The 2 New York local- ities shown in the figure are the Bronx River and are taken from Crocker (1957:70). Specific localities for New England have not been published and therefore I list the following 21 records by state. MAINE (1) Hancock Co., George’s Pond in Franklin, coll. by W. F. Reid, Jr., 29 Aug 1970. MASSACHUSETTS 7 collections made by Tom J. Andrews: (2) Worcester Co., Upton township, inlet to Pratt Pond, draining Dean Pond, near Upton-Hopkinton Rd., 6 June 1956. (3) Plymouth Co., Bridgewater township, Lake Nippenicket near Lake- side Drive and rt. 104, 25 Nov 1955. (4) Bristol Co., Dorchester Brook, 50 yds below Monte Pond in Brock- ton, 20 June 1953. (5) Bristol Co., Norton township, SW end of Norton Reservoir, 31 Aug 1957. (6&7) Barnstable Co., Mill Pond in Brewster-Harwich townships, 28 Sept 1958 and 25 Aug 1959. (8) Barnstable Co., Brewster township, Small’s Pond, 25 Aug 1959. 238 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 2 collections made by Bruce B. Collette: (9) Barnstable Co., Mills R. at rt. 28, 19 Sept 1957 (BBC 380). (10) Norfolk Co., Eagle Brook along rt. 140 between Foxboro and Wrent- ham, 21 Aug 1956 (BBC 247). 1 collection which I made: (11) Hampshire Co., Easthampton topographic sheet, Mill R. upstream from bridge at crossing of rt. 10, 1 mi SW of Northampton, Conn. R. drainage, 8 Aug 1953. This locality has been discovered indepen- dently by Douglas Smith of the Museum of Zoology, Univ. Mass., in collecting between 1972 and the present. RHODE ISLAND 2 collections made in Kent Co. by D. W. and R. A. Crocker on 27 Aug Ova: (12) trib. of Big R. at rt. 3, 0.3 mi S of Noosneck at Noosneck Hollow Bridge. (13) S arm of Flat R. Reservoir at Harkney Hill rd. crossing. 8 collections from the Dept. of Zoology, Univ. R.L.: (14) Kent Co., small pond in W Warwick, SW of junc. of London turnpike and rt. 3, coll. by H.J.C., 15 April 1963. (15) Kent Co., Breakhart Brook at NE edge of Arcadia Management Area in West Greenwich, coll. by Mallen, 24 April 1965. (16) Providence Co., stream at crossing of rt. 103, 0.5 mi E of Riverside, coll. by Brewster, no date. (17) Providence Co., N shore of Sneech Pond about 0.5 mi NE of Cum- berland Hill, coll. by P. Izra, 1 April 1962. (18) Providence Co., Olney Pond in Lincoln Woods State Park, coll. by Harting, 12 May 1956. (19) Providence Co., Clear R. near junc. with Wilson Reservoir in Bur- relville, coll. by D. J. Zinn, 22 April 1952. (20) Washington Co., Carolina Hatchery at Arcadia, coll. by Woods and Bishop, 20 July 1960. (21) Washington Co., stream in Tuckertown entering Worder Pond, coll. by J. Stedman, 1 May 1963. The single locality in Maine is so distant from those in Massachusetts that I regard it as an introduction by man, especially so in that 16 coastal col- lection sites, 6 of them in the general area of this locality, produced no VOLUME 92, NUMBER 2 239 crayfishes at all. The locality close to the Connecticut River in Massachu- setts results from collecting on 8 Aug 1953 (Mill R. upstream from bridge at crossing of Mass. rt. 10, 1 mi SW of Northampton). The continued pres- ence of P. a. acutus at this site is documented by 2 lots in the Museum of Zoology, University of Massachusetts, taken between 1972 and the present, reported to me by Douglas G. Smith (pers. comm.) of the Museum. The northern natural limit for this species in the east lies in the southern half of Massachusetts as may be seen in Fig. 3. Although there are no records from Connecticut, this state also probably contained and may yet be found to contain P. a. acutus. The distribution in New England as now known strongly suggests a movement into Cape Cod through Connecticut, possibly via marginal glacial lakes. Cambarus (Puncticambarus) robustus Girard The cluster of 5 New York localities (Fig. 3) is taken from Crocker (1957:82, Fig. 6). These lower Hudson River sites were then known to be connected with 12 others in the upper Hudson and upper Mohawk Rivers by only one collection: Schoharie Creek in Schoharie County (New York State Museum crayfish collection no. 2514, taken on 26 July 1934). This collection consists of only one very small female and the identification might be questioned, but in June 1971 I located C. robustus in quantity in Rens- selaerville, New York, in a tributary of Catskill Creek only 13 mi E of the Schoharie Creek site. Thus a population of C. robustus about half way between the southern and northern Hudson River drainage populations is assured. . On the basis of the distribution of C. robustus as we knew it when Barr and I were studying crayfish distributions in Ontario, it appeared that al- though the 5 southern New York collections could represent a relict pop- ulation, they might have been derived from introductions by man. There- fore, they were omitted from the general distribution of the species (Crocker and Barr, 1968:121, inset map in Fig. 82). Now, however, I am able to report C. robustus from 4 localities in Con- necticut (Fig. 3). A fifth locality is given by Bell (1971:14), ‘‘Bigelow Brook, Ashford [Windham Co.].’’ This and my localities 1 and 2 (below) are in the Thames R. system. My 4 localities are: (1) Fenton River, Tolland Co., Mansfield; collected by Mildred C. Ro- kowski, 15 April 1953. (2) Fenton River, Tolland Co., Gurleyville; collected by Univ. Connect- icut Field Zoology class, 20 July 1954. (3) West Branch Aspetuck River, Housatonic R. drainage, Litchfield Co., about 2 mi W of Northville; collected by D. W. and R. A. Crocker, 14 Aug 1971. 240 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON (4) Shepaug River, Housatonic R. drainage, Litchfield Co., Woodville; collected by D. W. and R. A. Crocker, 24 Aug 1971. There are thus 2 sets of sites, separated from each other by the Con- necticut River drainage system. If we assume that this gap is an artifact of collecting or a result of recent extinction from this system, then it now appears that the localities for C. robustus in the lower Hudson drainage are not introductions by man or relict populations, but are part of an east-west series of populations extending into New England. The total distribution of C. robustus is, in addition to the above, largely in portions of the states of Pennsylvania, Ohio, Michigan, Indiana, West Virginia, and Kentucky. It is known also from small parts of the states of Virginia, Tennessee, and North Carolina in the region where these three states meet (Crocker and Barr, 1968:121, inset map in Fig. 82; Hobbs, 1969: 134, Fig. 7; Hobbs, 1974b:22). C. robustus is absent from eastern West Virginia (Newcombe, 1929), Maryland (Meredith and Schwartz, 1959), east- ern Pennsylvania (Ortmann, 1906), New Jersey (Francois, 1959) and the Delaware and Susquehanna river systems in New York (Crocker, 1957:82). The current study has not found C. robustus in either Massachusetts or Vermont. It follows from the fact of the absence of C. robustus from these areas that its entry into Connecticut, the only New England state in which it is found, must have been at the western edge of the state via the lower Hudson River system. Thus, C. robustus is like P. a. acutus in having entered New England at its southwestern corner, but C. robustus entered from a northwesterly instead of a southerly direction. It seems reasonable to assume that C. robustus entered New England later than P. a. acutus since its movement eastward in New York must have been to the north of the Susquehanna and Delaware drainage systems. Such movement could not have occurred until recession of the glacial margin to that drainage border which lies about 125 mi N of the southernmost point at which P. a. acutus could have entered Connecticut. The validity of this assumption is strengthened by the extension of the range of C. robustus less far eastward than that of P. a. acutus. Cambarus (Cambarus) bartonii (Fabricius) Fig. 3 shows this species to be distributed in New England in association with the Appalachian mountain chain. This is in accord with Ortmann (1906:447) with whom both earlier and later students agree that, ‘‘We clearly see that its range follows the main strike of the Appalachian system, and . . . that ecologically this species is a form of the rapid and cool waters of the uplands and mountains, living preferably in small streams and even springs ....’’ Water temperatures for 5 August collections made in Mas- sachusetts and Vermont range from 14°C to 21°C with a mean of 17.7°C. VOLUME 92, NUMBER 2 24] The northernmost locality in Vermont, though not typical of the sites sam- pled in this survey, is not unusual for the species and is here described as an illustration of a species habitat unique among those from which I have collected New England crayfishes, and to provide an explanation for the distribution of C. bartonii along a mountain chain. The locality is Franklin County, township of Fletcher, about 4 mi S of Bakersfield on Vt. rt. 108 in a tributary to Black Creek. On 19 Aug 1970 the stream bed was dry at the road, but we followed it upstream on a 30° slope for about 0.5 mi until we reached a series of small pools. Here, search- ing for 45 min produced 1 male I, 2 males II, 1 female, and 10 immatures ranging from 7.0 to 16.5 mm carapace length. The water temperature was 15°C. Such habitats in gaps would allow crossing a divide overland and also they are susceptible to stream capture. The apparent rarity of C. bartonii probably is in part an artifact of sam- pling. Relatively few typical sites were visited, and also it is my impression that even in those localities where it exists it is less abundant than other species of crayfishes. Scattered observations of feeding activities suggest that C. bartonii is more consistently carnivorous than other New England crayfishes. This would place it higher on a pyramid of numbers and would suggest a lesser abundance. However, eastern New Hampshire and Mas- sachusetts as well as all of Connecticut and Rhode Island are coastal plain areas and their streams offer less suitable habitats for C. bartonii. Its ab- sence there must be due at least in part to this ecological factor. Hagen (1870:79) gives a seemingly anomalous locality record: *‘Aquarial garden, Boston.’’ Rathbun reports (1905:19), “‘C. bartoni introduced into brook at New Haven in 1880, but none have been seen there since.’’ Her species locality, Houlton, Maine, is the only record plotted in Fig. 3 which is taken from the literature. Clearly an inhabitant primarily of the Appalachian chain, C. bartonii has moved northward into Maine, Quebec, and New Brunswick (Faxon 1885:60-61). Norton (1909) summarizes the distributional picture in Maine as known at that time. In Ontario it has become distributed as far west as the east shore of Lake Superior, but is rare in the region between Lakes Erie and Huron. Crocker and Barr (1968:113-—115) document the above and describe the general distribution. Crocker (1957:84) has shown this subspe- cies to be widely distributed in New York, but rare in the streams entering the southern edge of Lake Ontario. It is present throughout Pennsylvania (Ortmann, 1906:381—382, 447, plate 43). In the mountains it reaches as far south as Georgia (Hobbs, 1974b:11). Orconectes propinquus (Girard) Fig. 4 shows that in New England O. propinquus is confined to the Lake Champlain and Hudson River drainage systems of Vermont and Massachu- 242 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON LEGEND BOUNDARIES NATIONAL & STATE =———»= = COUNTY WATERSHEDS SCALE OF MILES ~ t i ai \ ey Nees SOURCE: Traced (except drainage divides) from ‘ "THE NATIONAL SURVEY MAP OF NEW ENGLAND" The National Survey, Chester, Vt. 05143 Fig. 4. Species localities for Orconectes propinquus and O. rusticus in New England. O. propinquus = dark circles; O. rusticus = circles with lower half dark. VOLUME 92, NUMBER 2 243 setts with the exception of 4 localities: 1 just over the divide in the Con- necticut River drainage, 2 just over the divide in the Housatonic River drainage, and 1 about 60 stream miles downstream from the previous 2 localities in the Housatonic River. The 5 New York localities in Fig. 4 are additional to those reported by Crocker (1957:78). The distribution of this species is now fairly well known except for Que- bec. Crocker and Barr (1968:71—73) delimit the general distribution as the southeastern half of Ontario reaching as far north as James Bay, the Lake Ontario drainages in New York (upper Hudson and Mohawk Rivers should have been included), and the northern border of Ohio. It is a common species throughout Indiana, and is in upper and lower Michigan, Wisconsin, and eastern Iowa. O. propinquus apparently has entered New York and its limited area in New England by moving through glacial Lake Maumee and its subsequent stages, and by entering the St. Lawrence when it was formed. Orconectes limosus (Rafinesque) Localities plotted in Fig. 5 show this species in New England to be an inhabitant primarily of the coastal plain areas. The general distribution of this crayfish as previously known is northern Virginia, District of Columbia, eastern Pennsylvania, northwestern Maryland, western and northern New Jersey, Middlesex County in Connecticut, and Berkshire and Essex counties in Massachusetts (Rhoades, 1962:89). Crocker (1968:64) reports O. limosus to be in 3 adjacent drainages in Maine, ‘“‘Sebago Lake—Presump- scot River, Kennebec—Androscoggin Rivers, and Penobscot River (and the adjacent Orland River).’’ Aiken (1964) found O. limosus in New Hampshire in Newfound Lake (and Newfound River below the lake), and in Conway Lake. Osburn (1912:924) reported receiving a letter from A. E. Ortmann with the information that O. limosus had been introduced into a lake in East Hampton, Connecticut. Douglas Smith (pers. comm.) adds Barnstable Co. to the Massachusetts records for O. limosus with 2 collections in the Mu- seum of Zoology, University of Massachusetts: Orleans (no indication of body of water) and Brewster (Cliff Pond). Webster (1944:126) recorded the presence of O. limosus in Columbia Lake, Tolland Co., Connecticut. These also are coastal plain areas. Crocker and Barr (1968) did not find O. limosus in Ontario. Crocker (1957:78) recorded the species as inhabiting in New York the Susquehanna, Delaware, and lower Hudson Rivers. No unquestioned localities for this crayfish have been reported previously for the Great Lakes-Lake Cham- plain-St. Lawrence River drainages. Crocker (1957:78—80) discussed 4 ques- tionable localities. Fig. 5 shows localities for O. limosus at the N and S ends of Lake Champlain. Bell (1971:15) says it is common in shallow bays 244 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON ——$-—---—— ra 7 LEGEND BOUNDARIES ne NATIONAL & STATE == ——-— = COUNTY 9 WATERSHEDS = ---—---— SCALE OF MILES \ q = ath = lf 7 i i 1 eae Late i i 5 SOURCE: Traced (excep! drainage divides) from A yal W "THE NATIONAL SURVEY MAP OF NEW ENGLAND" $ The National Survey, Chester, Vt. OSI43 — tL = o = =oe5 fe Ae, = = 5 f To" 6r30" or —_ i Fig. 5. Species localities for Orconectes limosus in New England. VOLUME 92, NUMBER 2 245 of the lake and he gives 2 specific localities about midway along the length of its drainage area in Vermont: Shelbourne Pond and a pond in Winooski. At the time of my report on New York crayfishes (Crocker, 1957:78), O. limosus was not known in that state north of the mid-section of the Hudson River drainage, although the area was not sampled intensively. To my knowl- edge the western shores of Lake Champlain have not been searched for crayfishes subsequently, but I assume now that this species may be widely distributed in the Lake Champlain system. O. limosus has entered New England from the lower Hudson River drain- age system. Orconectes virilis (Hagen) Fig. 6 shows localities for this species in New England. In addition, 2 are included for Quebec (E shore, Three Sisters Island, Lake Memphramagog; small stream near Bedford). Six of the New York localities are repeated from Crocker (1958:71, Fig. 3), but the seventh (near the intersection of the state borders of New York, Vermont, and Massachusetts) is new. Crocker and Barr (1968:94—96) and Crocker (1957:72) give reference to studies from which the following picture of the general distribution of this species is drawn. O. virilis is the most widely distributed and abundant crayfish in Ontario. It ranges through the northern parts of Ohio, Indiana, Illinois, and Iowa. It occurs throughout the states of Michigan and Wisconsin. There are collections in NMNH from Atlantic Co., New Jersey and Summers Co., W. Virginia, and from several localities in Kansas and Missouri. It has not been found in Pennsylvania or Kentucky. Its southwestern limits are unclear, but several Montana localities are known. Camougis and Hichar (1959) studied O. virilis in Hobbs Brook Reservoir in Lincoln, Massachusetts. Aiken (1965) reports this species from Great East Lake, Province Lake and Townhouse Pond on the Maine-New Hamp- shire border, and from Lake Winnipesaukee. He records its absence from Newfound Lake, and Conway Lake. In addition (pers. comm., 19 Sept 1969) he has supplied me with the following list of lakes in New Hampshire from which he has taken O. virilis: Crystal Lovell Squam Wentworth Goose Mascoma Suncook Wickwas Kanasatka Merrymeeting Swains Winnisquam Little Squam Mirror Waukewan He reports the following lakes (or ponds) as lacking crayfish: Bow, Dan Hole Pond, Ossipee, Pine River Pond, Silver, Stinson. Douglas Smith (pers. comm.) reports a collection of O. virilis from the Connecticut River drainage of Massachusetts in the Museum of Zoology, University of Massachusetts: Hampshire Co., Mill River in Amherst. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 246 LEGEND BOUNDARIES NATIONAL & STATE —--:=---—-= COUNTY WATERSHEDS SCALE OF MILES i oo Cy k by SOURCE: Traced (except drainage divides) from "THE NATIONAL SURVEY MAP OF NEW ENGLAND" The National Survey, Chester, Vt. 05143 New England. . is in Species localities for Orconectes viril Fig. 6. VOLUME 92, NUMBER 2 247 I am not able to determine whether the patchwork distribution shown in Fig. 6 more nearly represents a natural or an artificial distribution, or in what proportions the agencies of man and nature have formed it. I have mentioned in an earlier section that transfers of this species within the area have occurred. The movement of O. virilis eastward toward New England appears to have been through Ontario in postglacial times. In Ontario at present it is the crayfish with the most widespread distribution (Crocker and Barr, 1968:95). Five scattered localities in the Hudson River drainage system in New York seem best explained as introductions in that an early colonization probably would have achieved a spread throughout the state. Crocker (1957:71) shows that this spread has not occurred and that O. virilis is located only at the western, northern, and eastern borders. O. virilis has been shown to be an aggressive species which when introduced can expand its territory to the disadvantage of native species (Bovjberg, 1961 and 1970; Schwartz et al., 1963). Hobbs (1974b:43) includes Maine in the natural range of this species. He records it as introduced into California, Maryland, Tennessee, and ‘‘.. . parts of New England... .”’ Orconectes immunis (Hagen) Fig. 7 shows localities for this species in New England. The following outline of its total distribution is taken from Crocker and Barr (1968:106— 107) where references are made to the supporting literature. O. immunis occurs throughout Indiana and Illinois. It is also in northwestern Tennessee, east-central Oklahoma, northeastern Kansas, the southern portion of Mich- igan, southeastern Ontario, northern New York, the southern half of Wis- consin, and southern Nebraska. Its greatest extent westward is in north- western Colorado. This species has not been found in Pennsylvania, New Jersey, or West Virginia. A few scattered records for New Hampshire and Massachusetts are given by Creaser (1933:16) and Faxon (1914:378). Aiken (1965) reports collecting this species in Newfound Lake in New Hampshire and he notes a collection of it in the Museum of Comparative Zoology, taken in Tilton, New Hamp- shire, in 1941. Douglas Smith (pers. comm.) reports a collection of O. im- munis from Great East Lake, Carrol Co., N.H., and 2 collections from Hampshire Co., Mass. These are in the Museum of Zoology, University of Massachusetts. In the section above on introduction and transfers I pointed out that Faxon (1914) gives considerable attention to the Massachusetts records. He comes to the conclusion that (p. 381), “*. . . Berkshire County is the eastern limit of the natural distribution of this species ... ,’’ but he notes, “‘for what it is worth. . . the Berkshire countrymen whom I have PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 248 LEGEND BOUNDARIES NATIONAL & STATE === == COUNTY WATERSHEDS SCALE OF MILES (see note) SOURCE: Traced (except drainage divides) from “THE NATIONAL SURVEY MAP OF NEW ENGLAND" The National Survey, Chester, Vt. O5143 Species localities for Orconectes immunis in New England. Fig. 7. VOLUME 92, NUMBER 2 249 questioned believe that crayfishes are a comparatively late addition to the fauna of the lakes [Pontoosuc Lake, Onota Lake, Lanesborough Pond, Goodrich’s Pond].”’ I am unable to offer with confidence a rationale for the distributional pattern of O. immunis in New England. One problem is that in my contacts with bait dealers in New York, I have found O. immunis to be the species most commonly propagated in artificial ponds. Forney (1956:6) recommends it as the only species of crayfish to try to culture. Some of the New England species localities for O. immunis may well have resulted from the activities of bait dealers and from their customers as well. This is just one aspect of the general problem of introductions and transfers discussed earlier. Considering the direction of movement of O. immunis toward New En- gland and its area or areas of entry, one can be somewhat more confident. Looking at maps of the distribution of this species in New York (Crocker, 1957:73) and Ontario (Crocker and Barr, 1968:107), this species is seen to be resident in many streams along the southern border of Lake Ontario, and well represented in Ontario in the area between Lake Erie and Lake Huron- Georgian Bay. Seven sites in the Susquehanna River drainage in New York are all close to the divide between it and the Lake Ontario watershed, and I consider them to be introductions or recent migrants over the divide south- ward. This picture suggests, considering also that O. immunis is not in Pennsylvania, that if this species moved eastward by natural means into New England, it did so at least in part through the area in New York where streams now drain northward into Lake Ontario. It could then have moved through the current Mohawk River valley and mid-portion of the Hudson River where populations of it now exist. Did O. immunis nove eastward through eastern Ontario and Quebec also, and enter New England from the north as well as from the west? Species localities for O. immunis are sparse in this area, possibly a result of inad- equate sampling. Crocker (1957:73) shows a locality on the northernmost border of New York and Crocker and Barr (1968:107) show 3 localities near the outlet of Lake Ontario and a cluster of 4 localities about 100 mi NW of this area. A collection of O. immunis in the Museum of Natural Sciences, National Museums of Canada, from a site SE of Bancroft, Haliburton Co. is roughly midway between these 2 sets of localities (Diana Laubitz, pers. comm.). Bousfield (1969) does not include O. immunis among the crayfishes of the Ottawa region. Figure 7 shows 6 localities in the northern half of Vermont. If this total of 14 species localities represents a natural distribu- tion, then either O. immunis has migrated eastward in Ontario via the Kirk- field or Ottawa outlets (Crocker, 1957:72) or it has reached these localities by spreading eastward and northward around the S and E sides of Lake Ontario through New York, having gained entry to that state perhaps in 250 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Lake Maumee time. Entry into New England could have occurred in several areas along the New York-Vermont and New York-Massachusetts borders. New State Records To the state records for crayfish species summarized from the previous literature on page 3, the current study adds the first specific locality infor- mation for several species as follows: Connecticut—Orconectes rusticus, introduced. It is still not certain that Cambarus bartonii is present and it is conjectured that Procambarus a. acutus is present. Maine—O. immunis and O. virilis and as introductions, O. obscurus, O. rusticus, and P. a. acutus. Massachusetts—P. a. acutus and as introductions, O. obscurus and O. rusticus. New Hampshire—O. rusticus (introduced). Rhode Island—O. immunis, O. limosus, O. virilis, and P. a acutus; P. clarkii (introduced). It seems unlikely that C. bartonii occurs here nat- urally if at all. Vermont—O. rusticus (introduced). Acknowledgments Financial support was provided by a research grant (GB-12287) from the National Science Foundation and by faculty research grants from Colby College and Skidmore College. Colleague friends have been generous in sharing their collecting time and in providing specimens as gifts or on loan: Tom Andrews and Douglas Smith (Univ. Mass.), Robert Chipman and Don- ald Zinn (Univ. R.I.), John Rankin (Univ. Conn.). Twenty-five collections from Connecticut were garnered in 1953 by Mildred Rokowski in the course of her study of the crayfishes of that state (M.S. thesis, Univ. Conn., 1954). The base map was traced and localities plotted by Christine Pheteplace Kloppenburg with permission from the National Survey, Chester, Vt. My son Ralph A. Crocker assisted on 2 collecting trips. Douglas Mathieu as- sisted with collecting in Maine. Horton H. Hobbs, Jr. (National Museum of Natural History) has been encouraging not only throughout the long ges- tation of this paper, but he has supplied materials, information, and advice during the almost 30 years in which I have been pleased to study crayfishes. References Cited Aiken, David E. 1965. Distribution and ecology of three species of crayfish from New Hamp- shire.—Amer. Midl. Nat. 73(1):240-244. Andrews, E. A. 1906a. Geographical distribution of crayfish.—Johns Hopkins Univ. Circular 25:100-103. VOLUME 92, NUMBER 2 251 . 1906b. The future of the crayfish industry.—Science 23:983—986. Bell, Ross T. 1971. Handbook of the Malacostraca of Vermont and neighboring regions.— Privately printed, pp. 1-65. Bouchard, Raymond W. 1975. Geology and ecology of crayfishes of the Cumberland Plateau and Cumberland Mountains, Kentucky, Virginia, Tennessee, Georgia and Alabama. Part II. The genera Fallicambarus and Cambarus.—Pages 585-605 in James W. Avault, Jr. (ed.), Freshwater crayfish, Papers from the Second International Symposium on Freshwater Crayfish. Baton Rouge, Louisiana: Louisiana State University Division of Continuing Education, 676 pp. Bousfield, E. L. 1969. Crayfishes of the Ottawa region.—Trail and Landscape 3(4):119-126. Bovbjerg, Richard. 1961. Mechanisms isolating two species of crayfish.—Amer. Zool. 1:345. 1970. Ecological isolation and competitive exclusion in two crayfish.—Ecology 51:225—236. Camougis, George and J. K. Hichar. 1959. Some studies on crayfish distribution in a small pond.—Amer. Midl. Nat. 62(1):227-231. Creaser, E. P. 1933. Descriptions of some new and poorly known species of North American crayfishes.—Occ. Pap. Mus. Zool., Univ. Mich. (275):1-21. Crocker, Denton W. 1957. The crayfishes of New York State.—Bull. N.Y. State Mus. Sci. Serv. (355): 1-97. . 1958. The crayfish Orconectes limosus (Rafinesque) in Maine.—Maine Field Natu- ralist 14(3):63—66. Crocker, Denton W. and David W. Barr. 1968. Handbook of the crayfishes of Ontario.— Univ. Toronto Press, xiii + 158 pp. Erichson, W. F. 1846. Ubersicht der Arten der Gattung Astacus.—Arch. f. Naturgesch. 12(1):86—103, 375-377 Fabricius, Johann C. 1798. Supplementum Entomologiae Systematicae.—Hafniae: Proft et Storch, 572 pp. Faxon, Walter. 1885a. Descriptions of new species of Cambarus; to which is added a syn- onymical list of the known species of Cambarus and Astacus.—Proc. Amer. Acad. Arts and Sci. 20:107-158. . 1885b. A revision of. the Astacidae. Part I. The genera Cambarus and Astacus.— Mem. Mus. Comp. Zool. 10(4):1—186. . 1914. Notes on the crayfishes in the United States National Museum and the Museum of Comparative Zoology.—Mem. Mus. Comp. Zool. 40(8):347—427, 13 plates. Forney, John. 1956. Raising crayfish for bait—N.Y. State Conservationist 10(6):6—7. Fowler, H. W. 1912. The Crustacea of New Jersey.—Ann. Rept. N.J. Mus. for 1911, Part IJ:29-650, 150 plates. Francois, Donald D. 1959. The crayfishes of New Jersey.—Ohio J. Sci. 59(2): 108-127. Girard, Charles. 1852. A revision of the North American Astaci, with observations on their habits and geographical distribution.—Proc. Acad. Nat. Sci. Philadelphia 6:87—91. Gould, A. A. 1841. Report on the Invertebrata of Massachusetts.—Cambridge, Mass. Hagen, Hermann A. 1870. Monograph of the North American Astacidae.—lIllus. Cat. Mus. Comp. Zool. Harvard 3:1-109. Hobbs, Horton H., Jr. 1942. A generic revision of the crayfishes of the subfamily Cambarinae with a description of a new genus and species.—Amer. Midl. Nat. 28(2):334—-357. . 1969. On the distribution and phylogeny of the crayfish genus Cambarus.—In Perry C. Holt (ed.), The distributional history of the biota of the southern Appalachians, Part I: Invertebrates. Va. Polytechnic Inst., Res. Div. Monograph, 1:93-178. . 1972. The subgenera of the crayfish genus Procambarus (Decapoda: Astacidae).— Smithson. Contrib. Zool. (117):1-22. . 1974a. Synopsis of the families and genera of crayfishes (Crustacea: Decapoda).— Smithson. Contrib. Zool. (164):111 + 32 pp. 252 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON . 1974b. A checklist of the North and Middle American crayfishes (Decapoda: Astacidae and Cambaridae).—Smithson. Contrib. Zool. (166):iii + 161 pp. Meredith, W. G. and F. J. Schwartz. 1959. The crayfishes of Maryland.—Maryland Tidewater News 15(1), supplement No. 12:1-2. Momot, Walter T., Howard Gowing, and Patricia Jones. 1978. The dynamics of crayfish and their role in ecosystems.—Amer. Midl. Nat. 99(1):10-35. Newcombe, Curtiss L. 1929. The crayfishes of West Virginia.—Ohio Jour. Sci. 29(6):267— 288. Norton, Arthur H. 1909. Some aquatic and terrestrial crustaceans of the State of Maine.— Proc. Portland Soc. Nat. Hist. 2:245—256. Ortmann, Arnold E. 1905. Procambarus, a new subgenus of the genus Cambarus.—Ann. Carnegie Mus. 3(3):435—442. . 1906. The crayfishes of the state of Pennsylvania.—Mem. Carnegie Mus. 2(10):343- 533. Osburn, Raymond C. 1912. The crayfish.—Bull. N.Y. Zool. Soc. 16(54):921-927. Rafinesque, C. S. 1817. Synopsis of four new genera and ten new species of Crustacea, found in the United States—Amer. Monthly Mag. and Crit. Rev. 2:40—43. Rathbun, Mary J. 1905. Fauna of New England. 5. List of the Crustacea.—Occ. Pap. Boston Soc. Nat. Hist. 7:1-117. Rhoades, Rendell. 1962. The evolution of crayfishes of the genus Orconectes section Limo- sus.—Ohio Jour. Sci. 62(2):65—96. Sanford, S. N. F. 1932. Marine invertebrates, in Museum Department Reports.—Bull. Boston Soc. Nat. Hist. (64):18. . 1936. Marine invertebrates, in Museum Notes.—Bull. Boston Soc. Nat. Hist. (78):11. Spohrer, Mary L., Jon L. Williams, and James W. Avault, Jr. 1975. A selected bibliography of the red swamp crayfish, Procambarus clarkii (Girard) and the white river crayfish, Procambarus acutus acutus (Girard).—Pages 637-661 in James W. Avault, Jr. (ed.), Freshwater crayfish, Papers from the Second International Symposium on Freshwater Crayfish. Baton Rouge, Louisiana: Louisiana State University, Division of Continuing Education, 676 pp. Schwartz, Frank J., Robert Rubelmann and James Allison. 1963. Ecological population ex- pansion of the introduced crayfish, Orconectes virilis.—Ohio Jour. Sci. 63(6):266—273. Thompson, Zadock. 1842. A history of Vermont, natural, civil, and statistical, with a new map of the State and two hundred engravings.—Burlington, Chauncey Goodrich. Webster, Dwight A. 1944. Notes on the food of smallmouth bass (Micropterus dolomieu) in Columbia Lake, Connecticut.—Copeia 1944(2):125—126. Williamson, William D. 1832. The history of the State of Maine from its first discovery, A.D. 1602, to the separation A.D. 1820, inclusive.—Vol. 1, xii + 660 pp; vol. 2, vii + 714 pp. Glazier, Masters & Co., Hallowell, Maine. Biology Department, Skidmore College, Saratoga Springs, N.Y. 12866. PROC. BIOL. SOC. WASH. 92(2), 1979, pp. 253-271 THE MARINE ISOPOD CRUSTACEANS OF THE GULF OF CALIFORNIA IT. IDOTEIDAE: NEW GENUS AND SPECIES, RANGE EXTENSIONS, AND COMMENTS ON EVOLUTION AND TAXONOMY WITHIN THE FAMILY Richard C. Brusca and Barry R. Wallerstein Abstract.—Three new records of Idoteidae (Isopoda; Valvifera) in the Gulf of California are presented, bringing the total known idoteid species of this region (the Cortez Province of Briggs, 1974) to 11. One new genus and species (Parasymmerus annamaryae) and one formerly described species (Cleantioides occidentalis) are fully described and figured. A key to the Idoteidae of the Gulf of California is presented. Parasymmerus is a highly derived taxon that appears to have evolved from Eusymmerus (or a common ancestor). Edotea apparently evolved as a tropical component of the Transisthmian Track fauna, but was quickly excluded to the colder temperate waters north and south of the American tropics where it exists today. Synidotea harfordi is-a widely ranging species, whose occurrence in the Cortez Province is uncommon. Cleantioides occidentalis is a widely ranging tropical/subtropical species, extending through 4 zoogeographic provinces. In light of recent morphological discoveries (in this paper and elsewhere) it iS apparent that the genera Cleantis, Cleantioides, and Zenobiana are in need of reexamination. Taxonomic characters of the Idoteidae are discussed and the use of “‘dor- sally visible coxal plates’’ or ‘‘dorsally visible coxal plate sutures’’ is seen to be largely unreliable. Presence or absence of dorsal coxal plates, and their general morphology, are important characters in the Idoteidae, and workers are urged to describe these structures more fully, and to distinguish between ‘‘dorsal coxal plates’’ and ‘‘ventral coxal plates.’ Brusca and Wallerstein (1977) reported the presence of 8 species of idoteid isopods from the Gulf of California. We report herein the presence of 3 additional species, one of which represents a new genus and species. The materials upon which this study is based are derived primarily from 3 sources: recently discovered collections of the late E. Yale Dawson from Western Mexico; material from Scripps Institution of Oceanography; and specimens collected on an expedition to tropical west Mexico by the au- thors, accompanied by P. Pepe and A. M. Mackey, in the summer of 1976. 254 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Genera of the family Idoteidae are, at the present time, distinguished primarily by 4 characters: development of the pleonal somites; arrangement of the (dorsal) coxal plates; number of flagellar articles in antenna 2; and number of articles in the maxillipedal palp. A recent paper by Menzies and Miller (1972) reviews the subfamily Idoteinae, to which most of the Idotei- dae genera belong, and the criteria and terminology used in the present paper are largely those used by these workers (see also Menzies, 1950). The following abbreviations are used: AHF, Allan Hancock Foundation; USNM, National Museum of Natural History; NMC, National Museums of Canada, Ottawa; AMNH, American Museum of Natural History (New York). Parasymmerus, new genus Etymology.—The generic name is indicative of the close similarity to the genus Eusymmerus; gender masculine. Type-species.—The type and only known species of the genus is described below, Parasymmerus annamaryae 0. sp. Diagnosis.—Pleon comprised of a single segment, with one pair of small, partial, anterolateral sutures. Coxae of pereonites II to IV simple rings, not forming dorsal coxal plates; coxae of pereonites V and VI free, expanded anterolaterally into small dorsal coxal plates (visible only in ventrolateral aspect); coxae of pereonite VII free, plates visible in dorsal aspect. Antenna 1 much shorter than antenna 2. Antenna 2 of 6 articles, the sixth being the single, large, flagellar article. Maxillipedal palp of 3 articles. Cephalon with- out a dorsal hump. Affinities.—Parasymmerus closely resembles the genera Eusymmerus and Edotea. It is similar to Eusymmerus in the following respects: general body symmetry; relative lengths of antennae 1| and 2; pleonal characteristics; and possession of a single, large, flagellar article on antenna 2. Parasym- merus differs from Eusymmerus in possessing only 3 articles on the max- illipedal palp, lacking a cephalic hump, and in having a dorsal coxal plate (albeit small) on pereonite V. We have found that even in very young in- dividuals of Eusymmerus (less than 3.0 mm in length) the maxillipedal palp is always composed of 4 articles, never 3. Parasymmerus shares with Edo- tea the pleonal characteristics and a maxillipedal palp of 3 articles. It differs from Edotea in the possession of free dorsal coxal plates on pereonites V— VII, and in having a large, uniarticulate, clavate flagellum on the second antennae (see Table 1). Menzies and Miller (1972) put forth the hypothesis that there exists, in the subfamily Idoteinae, a phylogenetic trend towards increasing degrees of fusion of certain body parts—notably the pleonites, dorsal coxal plates, articles of the maxillipedal palp, and possibly the flagellar articles of the second antennae. What has in the past, however, been interpreted as fusion VOLUME 92, NUMBER 2 255 Table 1. Comparison of Some Generic Characters of Eusymmerus, Parasymmerus, and Edotea (family Idoteidae; subfamily Idoteinae). Number Number Pleonal articles articles sutures in fla- in max- Number pereonites (complete + gellum illipedal with dorsal coxal partial of 2nd Genus palp plates* sutures) antennatt Eusymmerus 4 2 (unfused) 0+ 1 1 Parasymmerus 3 3 (unfused) 0+ 1 1 Edotea 3 3 (fused or partially fused) 0+ IT 1-4 * This refers to the actual number of dorsal coxal plates present on pereonites II through VII (whether fused or not with their respective pereonite), not just those which are visible in the dorsal view alone (see text). + In some species of Edotea grooves may extend across the dorsum of the pleon (e.g. Edotea transversa). These grooves are not true pleonite articulations, but appear to be remnants of the embryonic fusion of the pleonal somites. ++ There appears to be two distinctly different types of reduction that has occurred on the flagellum of antenna 2 of the Idoteinae. In most genera possessing a uniarticulate flagellum (including Eusymmerus and Parasymmerus) the single article is a large, clavate structure, suggesting possible fusion of the flagellar articles. In Edotea, on the other hand, the flagellar articles are simply reduced in size and number, being minute vestiges borne upon the peduncle, with no suggestion of fusion having taken place. of the dorsal coxal plates to the pereonites appears to be in error in many cases. Our observations indicate that many assumptions of coxal plate fu- sion can be more correctly interpreted as coxal plate reduction or loss (see discussion below). This phylogenetic trend in the Idoteinae is best seen when the genera are arranged in a series of increasing fusion of pleonites and reduction in maxillipedal palp articles (see table 1, Menzies and Miller, 1972:2). We agree that these data rather convincingly suggest that the more primitive genera in the subfamily may be distinguished by a pleon comprised of 2 to 4 complete pleonites (plus | to 2 additional, partly fused pleonites, indicated by incomplete lateral suture lines), and a maxillipedal palp of 4 to 5 articles. The genera distinguished by these plesiomorphic character states are: Zenobiana, Cleantis, Cleantioides, Idotea, Cleantiella, Engidotea, and the extinct Proidotea. Genera possessing the presumably most derived, or apomorphic character states (all pleonites fused into a single piece, with no or a Single pair of partial lateral suture lines; maxillipedal palp of 3—4 articles) are: Moplisa, Colidotea, Synidotea, Synisoma, Erichsonella, Eusymmerus, Parasymmerus, and Edotea.! Parasymmerus is one of the most derived taxa in the Idoteinae, in having only a single partial pleonal suture and a maxillipedal palp of 3 articles. If a decreased number of flagellar articles in 1 Note that Menzies and Miller’s (op. cit.) table incorrectly states that Synisoma has 2-3 partial pleonal suture lines (it actually has only 1), and that Edotea has a second antennal flagellum of a single article (it actually has 1-4 rudimentary flagellar articles). 256 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON antenna 2 is also considered an apomorphic attribute, Parasymmerus ap- pears as a highly derived taxon indeed in its reduction to a single flagellar article. If the characters of Menzies and Miller (above) are used, the most recently derived genera of Idoteinae are seen to be members of the New World fauna, particularly the New World tropics, Parasymmerus apparently being derived from Eusymmerus or a common ancestor. These two genera, in turn, appear to be at or near the apex of a lineage that also produced Er- ichsonella, Colidotea, Edotea, and possibly the unusual southwest Atlantic genus Moplisa. A geographic trend is also apparent. Eusymmerus and Par- asymmerus are monotypic genera Known only from the tropical east Pacific. Edotea, on the other hand, is a polytypic New World genus, comprised of 5 species known from the temperate Pacific Southern Hemisphere, and 2 from the temperate Northern Hemisphere (E. sublittoralis [Pacific] and E. triloba [Atlantic]).” In order for this distributional pattern to exist one must assume, if the above phylogenetic hypothesis is accepted, that the genus Edotea evolved as part of the Transisthmian Track fauna (see Croizat, Nel- son, and Rosen, 1974) of the Central American waterway. Some early ances- tor(s) of the west Atlantic species must have once flourished in the tropical — Caribbean region, later to become extinct (at least locally), leaving as its descendant the single temperate northwest Atlantic species Edotea triloba (Say). In the northeast Pacific only Edotea sublittoralis persists today— restricted to temperate offshore benthic waters, in subtidal depths to 64 m (Brusca et al., ms.). One may speculate that competition in the intertidal and shallow subtidal regions of the northeast Pacific, at the time of E. sublittoralis’ (or its ancestor’s) early colonization efforts was made severe by the presence of the already more speciose shallow-water genera Jdotea, Synidotea, and Celidotea (also possibly Cleantis and Cleantioides). Thus, we find that Edotea, although apparently evolving in the New World tropics, was promptly excluded from this region, its modern species being successful only in the temperate latitudes of North and South America. For a review of the distribution of the northeast Pacific Idoteidae see Brusca and Wal- lerstein, 1979. Parasymmerus annamaryae, new species Figs. 1-2 Etymology.—tThe specific name is the genitive singular of Anna Mary [Mackey]; it is named in Miss Mackey’s honor for her assistance on the expedition during which this new species was collected, as well as her col- lection of a great deal of other material upon which this and other isopod studies have been based. 2 Edotea acuta and Edotea montosa were synonymized with E. triloba by Wallace (1919). VOLUME 92, NUMBER 2 257 Fig. 1. Parasymmerus annamaryae. A, Dorsal view; B, Maxilliped; C, Uropod; D, Left mandible; E, Lacina mobilis of left mandible; F, Maxilla 1, endopod; G, Maxilla 1, exopod; H, Maxilla 2; I, Antenna 2; J, Antenna 1. 258 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Holotype.—AHEF 765; male. Type-locality.—Mexico, Colima, Manzanillo, Bahia Audiencia, north shore of bay, approximately 1 m below high tide mark in algae (Choonos- pora minima and Dermonema frappieri), dead barnacle shells (Tetraclita sp.), and mussel beds. Water (surf) temperature 30°C; air temperature (early afternoon) 27°C; overcast; light rain; 3-4 August 1977; A. M. Mackey, R. C. Brusca, B. Wallerstein, and P. Pepe (collectors). Allotype.—AHEF 765a; female. From same sample as holotype. Paratopotypes.—28 specimens; same sample as holotype. Deposited: AHF, USNM, NMC, AMNH. Paratypes.—1 nonovigerous female; Mexico, Sinaloa, Mazatlan; small ‘‘reef’’? 2 mi. north of town; 8 December 1946; E. Yale Dawson. Deposited AHF. Description.—Head at least 1.75 times wider than long, sides strongly produced laterally (Fig. la); supra-antennal line strongly concave; frontal process wide and truncate, extended almost to anterior margin of frontal lamina 1; frontal lamina 1 medially produced, wider than frontal process; frontal lamina 2 broadly rounded, visible in dorsal aspect. Eyes transversely elongate and wide, rarely subovoid. Antenna 1 short, at best reaching article 3 of antenna 2, and of 4 articles, the fourth being the single flagellar article bearing terminal esthetascs (Fig. 1j). Antenna 2 of 6 articles, the sixth being the single, large, flagellar article bearing short setae (Fig. 11). Maxillipedal palp of 3 articles, distal 2 articles with simple setae; endite with 1 coupling hook, 2 to 4 terminal spines, and 1 to 6 plumose setae along terminal margin (Fig. 1b). Maxilla 1 endopod with 2 to 3 long setae; exopod with 13 spines and 2 robust, simple setae (Fig. If,g). Maxilla 2 trilobate, endopod heavily setose, with 13-14 long setae, the outermost being plumose; inner lobe of exopod with 4 to 5 comb setae along distal margin; outer lobe of exopod with 4 to 6 comb setae (Fig. 1h). Mandible with 5-cuspate incisor process, robust molar process, and lacina mobilis of 3 cusps and a complex setation as figured; lacina mobilis of right and left mandibles subequal, although slight differences in setation do occur (Figs. 1d,e). Pereon broad, 1.7 to 2.1 times longer than pleon; lateral margins subpar- allel or slightly convex. Coxae of pereonites II to IV reduced, not forming dorsal coxal plates; dorsal coxal plates of pereonites V and VI very small, anteriolaterally directed, and visible only in ventrolateral aspect; dorsal cox- al plates of pereonite VII larger, triangular, posteriorly direct, with dorsally visible sutures (Fig. la). Pereopods 1 to 7 slender, ambulatory; dactyl with accessory claw. Pereopod 1 with comb setae on inner surface of propus, and weak setation on ventral margin (Fig. 2f). Pereopods 2—7 subsimilar, without comb setae but with weak setation on ventral margin (Fig. 2g). Pleotelson with posterolateral angles obtuse, tapering posteriorly to trun- cate tip. Uropod uniramous, with single, large, plumose seta on lateral distal VOLUME 92, NUMBER 2 259 Fig. 2. Parasymmerus annamaryae. A, First pleopod; B, Second pleopod; C, Third pleo- pod; D, Fourth pleopod; E, Fifth pleopod; F, First pereopod; G, Fifth pereopod. 260 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON angle of basis (Fig. 1c). Pleopods all bilamellar. Pleopod 1 strongly setose; lower lamella with plumose setae on medial margin; upper lamella with plumose setae on lateral margin and apex (Fig. 2a). Pleopod 2 with upper lamella subsimilar to that of pleopod 1; lower lamella with a few plumose setae on apex; appendix masculina of male extended beyond tip of lamella and acuminate, with tracts of minute spines distally (Fig. 2b). Pleopods 3 to 5 with simple setae (Figs. 2c,d,e). Distribution.—Although we have examined extensive collections from throughout northwestern Mexico, particularly the Gulf of California, these are the only two localities from which this isopod has been recorded. Par- asymmerus annamaryae is a tropical species and its range is expected to be extended considerably southward when more intensive collecting is accom- plished on southern Mexico and Central American rocky shores. Synidotea harfordi Benedict 1897 Fig. 3 Idotea marmorata Harford, 1877:117. Synidotea harfordi Benedict, 1897:402; Richardson, 1899a:849; Richardson, 1905:387; Gurjanova, 1936:163; Johnson and Snook, 1955:290; Schultz, 1969:67; Menzies and Miller, 1972:16; Miller (in Smith and Carlton), 1975:288, 289, 306. Diagnosis.—(After Menzies and Miller, 1972) Cephalon without preocular horns; frontal margin transverse or slightly convex, with no median emar- gination. Dorsal surface of pereon smooth, lacking rugae, tubercles, or scales (under light microscope); pereonites 1 to 3 with lateral margins evenly rounded; borders of pereonites 4 to 7 straight. Pleotelson about one-fourth longer than its greatest width, posterior border medially excavate. Appendix masculinum straight, apex bluntly pointed, lateral margin spined (Fig. 3a; for additional figures see Menzies and Miller, 1972). New records.—Mexico, Golfo de California, Baja California Sur (east coast), Punta Chivato, about 20 mi. south Santa Rosalia; subtidal; 12 August 1976; A. M. Mackey.—Mexico, Golfo de California, Sonora, Guaymas, Estero Soldado; near entrance to Estero; 20 April 1973, from University of Arizona Ichthyology Collection, UA 73-43. Distribution.—Although formerly known from Morro Bay, California, to Magdalena Bay on the west coast of Baja California Sur, this discontinuous range extension into the Gulf of California is not unexpected, and follows a pattern seen in at least two other members of the family Idoteidae (Brusca and Wallerstein, 1977): Idotea (Idotea) urotoma Stimpson and Idotea (Pen- tidotea) aculeata Stafford. Synidotea harfordi is hereby established as the southernmost ranging species of this principally cold water genus in North America. It is the only intertidal species in the genus known to occur in VOLUME 92, NUMBER 2 261 Fig. 3. Synidotea harfordi. A, Dorsal view; B, Maxilla 2 (from Guaymas, Sonora, Mexico). southern California and Baja California, and the only species of Synidotea known to occur in the Gulf of California. Remarks .—Although the specimen from Punta Chivato is not unusual in any regard, the specimen from Guaymas differs from the normal morphol- ogy in having a greatly enlarged outer lobe on the exopod of the second maxilla (Fig. 3b). Cleantioides occidentalis (Richardson 1899) Figs. 4—5 Cleantis occidentalis Richardson, 1899a:850; Richardson, 1899b:270; Rich- ardson, 1905:406; Richardson, 1912:28; Tattersall, 1921:426; Nierstrasz, 1941:265; Menzies, 1962:95; Menzies and Frankenberg, 1966:23; Schultz, 1969:83. Cleantioides occidentalis: Kensley and Kaufman, 1978:658. Diagnosis.—Body parallel sided, nearly 6 times longer than wide. Supra- antennal line with a small median emargination; apex of frontal process broadly rounded; frontal lamina 1 and 2 visible in dorsal aspect; lamina 1 262 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 4. Cleantioides occidentalis. A, Dorsal view; B, Maxilliped; C, Uropod; D, Maxilla 2; E, Left mandible; F, Maxilla 1; G, Antenna 1; H, Antenna 2. VOLUME 92, NUMBER 2 263 produced medially; lamina 2 truncate. Maxillipedal palp of 4 articles, endite with 2—4 coupling hooks. Lateral margins of body with dense tufts of plu- mose setae. Pleotelson with a pair of dorsal humps; uropods uniramous; appendix masculinum of male arises medially on endopod of pleopod 2. Pereopod IV greatly reduced and non-ambulatory. Description.—Body parallel sided, nearly 6 times longer than wide. Head wider than long; supra-antennal line with a small median emargination or notch; frontal process small, rounded apically, not extended to anterior margin of frontal lamina 1; frontal lamina 1 produced medially, extended almost to anterior margin of lamina 2; frontal lamina 2 wide and truncate. Eyes transversely (dorsoventrally) elongate, medial portion produced pos- teriorly (Fig. 4a). Antenna 1 of 4 articles, the fourth being the single flagellar article bearing terminal esthetascs (Fig. 4g). Antenna 2 of 6 articles, the sixth being the single flagellar article; all articles with 2 tracks of setae along ventral margin; articles 2—4 with scalloped distal margins (Fig. 4h). Maxil- lipedal palp of 4 articles, distal 3 with simple setae; endite with 2—4 coupling hooks, numerous plumose setae and spines along terminal margin (Fig. 4b). Endopod of maxilla 1 with 3 plumose setae; exopod with 12 spines and 3 plumose setae (Fig. 4f). Maxilla 2 trilobate, endopod heavily setose, with at least 14 setae, the outermost being plumose; inner lobe of exopod with 6 to 9 comb setae; outer lobe of exopod with 8 to 12 comb setae (Fig. 4d). Left mandible with 4-cuspate incisor, robust molar processes, and lacina mobilis of 3 cusps and complex setation (Fig. 4e); incisor and lacina mobilis of right mandible somewhat reduced in size. Pereon 2 to 3.2 times longer than pleon; lateral margins parallel and heavi- ly setose (Fig. 4a). Free dorsal coxal plates present on pereonites II to VII; those of V to VII occupying entire lateral margin, posterior angles acute; II and III occupying % lateral margin; IV occupying % to % lateral margin. Pereopods 1 to 3 slender, ambulatory, increasing in size posteriorly; per- eopod 4 greatly reduced and non-ambulatory, row of spines along terminal margin of last 4 articles, basis as long as 5 distal articles together (Fig. 5h); pereopods 5 to 7 increasing in size posteriorly (Fig. 51). All pereopods biun- gulate. Pereonites I-V, of female, bear oostegites. Pleon composed of 4 segments (including the pleotelson) plus one pair of partial sutures. Apex of pleotelson broadly rounded, with a pair of elevated, submedian, dorsal humps (Fig. 4a). Uropods uniramous, medial margins covered by dense setae, distolateral angle with single, large, plumose seta; distal margin with about 12 large, plumose setae (Fig. 4c). Pleopods all with plumose setae on outer margin of lamellae and on basis (Figs. 5a—f); pleo- pods 3-5 considerably less setose than 1-2 (Figs. 5d—f); marginal setae of exopods of pleopods 3 and 5 arise in 2 distinct tracks, one on the upper and one on the lower surface (Figs. 5d,f). Basis of pleopod 3 with distinct setose lobe (Fig. Sd). Appendix masculinum of male second pleopod arises not from basis, but medially from endopod (Fig. 5c). 264 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON S SN == SS Vs Vs WZ Ss SRO SSS SS VOLUME 92, NUMBER 2 265 New records.—Mexico, Sonora, SW of Punta Penasco; 12 m; 5-6 April 1960; from Scripps cruise, SB 25b; A. Flechsig.—Mexico, Baja California Sur, Santa Maria Bay (24°43’N 112°14'W); 35-50 m; 19 Jan. 1940; **Velero IlI,’’ station No. 1031—40.—Mexico, Michoacan, Punta Lizardo (18°6.7'N 102°56’W); 40’ otter trawl, 20-25 m; 4 April 1974; from Scripps cruise, MV 73-I-22; C. Hubbs and S. Luke.—Guatemala, off San Jose lighthouse (13°52'N 91°10’ W); 15-22 m; black sand; 11 Jan. 1930; ‘‘Velero III,”’ station No. 77-38.—Costa Rica, Gulf of Dulce (8°23’N 83°16'’W); 20—44 m; coarse sand; 26 March 1939; **Velero III,’ station No. 939-39.—Ecuador, off Cape San Francisco (0°37’N 80°0’ W); 30 m; mud and rock; 23 Feb. 1938; ‘‘Velero III,’ station No. 850-38.—Ecuador, Galapagos Islands, Charles Island, Black Beach (1°16’S 90°29’ W); rocky intertidal; 18 Jan. 1934; ‘‘Velero III,”’ station No. 163-34. Distribution.—Although this species was previously known only from the type-locality (Magdalena Bay, Baja California Sur), and Culebra Is., Pana- ma, our studies have revealed it to be a widely ranging tropical-subtropical species occurring through 4 zoogeographic provinces: the Cortez, Mexican, Panamanean, and Galapagos (Briggs, 1974; Brusca and Wallerstein, 1979). Collection data indicate that C. occidentalis has a preference for shallow, subtidal, sandy habitats (records are from the littoral region to 50 m). In addition, two of the collections were made on black sand substrates. Kens- ley and Kaufman (1978) report a salinity range of 26—-33%c in its habitat at Culebra Island. Remarks.—Richardson’s (1899a, 1899b) description of Cleantioides oc- cidentalis was based ona single individual, and was subsequently reproduced in her 1905 monograph. In neither of these treatments was this species adequately figured. Kensley and Kaufman’s (1978) redescription was, in turn, based upon 3 ovigerous females and 5 immature females, all from the same locality (Culebra Island, at the west entrance of the Panama Canal). Again, many of the appendages were not figured or described, and for these reasons an entirely new description and new figures have been prepared, based on males, females, and juveniles from west Mexico and Central America. Tattersall (1921) described the state of confusion that exists in the genera Cleantis and Zenobiana. Little has occurred in the subsequent 60 years to rectify this situation, and now Cleantioides Kensley and Kaufman (1978) must be included in this problem. These three closely related genera are in <— Fig. 5. Cleantioides occidentalis. A, First pleopod; B, Second pleopod, female; C, Second pleopod, male (setation not figured); D, Third pleopod; E, Fourth pleopod; F, Fifth pleopod; G, First pereopod; H, Fourth pereopod; I, Seventh pereopod. 266 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON need of overall reexamination, although Kensley and Kaufman did ‘‘pro- visionally’’ (sic) separate them based on examination of type species. They point out that Cleantis planicauda would seem to belong to the newly erect- ed genus Cleantioides, yet C. planicauda possesses a maxillipedal palp of 5 articles, rather than 4 as they have defined Cleantioides. If C. planicauda is indeed to be considered a Cleantioides, then the definition of this genus must be expanded to accomodate those species with a maxillipedal palp of 5 articles, and in doing so reducing the number of differences between Cleantis and Cleantioides to a single feature, a biramous vs. uniramous uropod. The only other genus in the Idoteinae to show variation in this maxillipedal character state is Idotea, but as Menzies and Waidzunas (1948) and Menzies (1950) have shown, species with a palp of 5 articles develop the fifth, small terminal article as they mature to adulthood. It was on this basis that Menzies reduced the genus Pentidotea to a subgenus of Idotea, and the number of articles in the maxillipedal palp remains the only character distinguishing these two subgenera. The variation in palp article number between C. occidentalis and C. planicauda, however, follows quite a dif- ferent pattern. Regardless of whether the palp is comprised of 4 or 5 articles, the small terminal article is always present, the additional article in C. planicauda apparently arising in the middle of the palp, between articles 2 and 3. In having only females to examine, Kensley and Kaufman (1978) were unaware of another important feature of C. occidentalis; in males the ap- pendix masculina of the second pleopod arises not from the basis, or even proximally on the endopod, but from the medial margin of the endopod (Fig. 5c). The significance of the unusual positioning of this structure is difficult to appraise, as other workers have failed to describe or figure the pleopods for other Cleantis species. Is this unusual configuration of the appendix masculina associated with a lineage in which the maxillipedal palp bears 4, rather than 5, articles? Due to the poor state of knowledge concerning the genera Cleantis, Cleantioides, and Zenobiana, the phyletic relationships of C. occidentalis are impossible to assess at this time.? Although C. occidentalis and C. planicauda are very similar in many regards, and are the only species of Cleantioides (or Cleantis) known from North America, they differ in several fundamental ways and cannot be considered twin or gemminate species. Further, we have a single record of C. planicauda from Oaxaca, Mexico, establishing this species as a probable amphi-American form, sympatric with C. occidentalis in the west American tropics. 3 Note that Menzies (1962) mistakenly stated that he had shown Cleantis occidentalis to be a juvenile of [dotea urotoma (Menzies, 1950). In fact, Menzies (op. cit.) synonymized C. heathii (not C. occidentalis) with I. urotoma. VOLUME 92, NUMBER 2 267 Remarks on the Coxal Plates and Their Use in Idoteid Systematics The following discussion uses a broad interpretation of Moore and McCormick’s (1969) definition of a coxal plate: the lateral expansion of a pereopodal coxa joined broadly to the lateral margin of the tergite. As Shep- pard (1957) has pointed out, however, in many valviferans the coxae may also be expanded ventrally, to form ventromedial plates (e.g. Edotea and Synidotea). These ventral plates are not, however, the structures upon which valviferan genera are traditionally diagnosed and separated. As Shep- pard (1957) and Bowman (pers. comm.) have noted, perhaps the time has come to begin discriminating between these two types of coxal structures by referring to them as dorsal coxal plates and ventral coxal plates (as we have done in this paper). Menzies and Miller (1972) gave considerable weight to the degree of sup- posed fusion of the dorsal coxal plates with their respective pereonites. Unfortunately, their table is based principally upon whether or not the plates (‘‘epimeres’’ sic) and sutures are visible in the dorsal aspect, NOT whether or not the coxae are actually expanded to form true dorsal coxal plates. We have found considerable variability in this character in several idoteid genera. In at least one species of Erichsonella (E. pseudoculata Boone) coxal plates are dorsally visible only on pereonites V to VII, but are actually present on II to VII. In our studies of Colidotea (Brusca and Wallerstein, 1977; Brusca et al., ms.) we have found that, in C. findleyi Brusca and Wallerstein, dorsal coxal plates are actually present on pereon- ites II to VII, although not dorsally visible on ANY segments (all bear distinct suture lines); in C. edmondsoni Miller coxal plates are dorsally visible on pereonites IV to VII; and in C. rostrata (Benedict) they are present on V to VII but dorsally visible on V to VII or VI to VII. In many idoteids the dorsal coxal plates may be quite small, particularly on the anterior pereonites, and visible only in ventrolateral view, with strong illumination. In Parasymmerus, for example, coxae V are only slightly ex- panded, VI somewhat more expanded, and VII fully expanded and visible in the dorsal aspect. This gradually increasing expansion (or reduction) in coxal size makes interpretation of coxal plate presence difficult. In other genera and species the trend towards reduction in the size of certain coxae is such that no true dorsal coxal plates are even formed. This reduction has mistakenly been interpreted as a fusion of the coxal plates to the terga of their respective pereonites by most workers (see, for example, Sheppard’s [1957] clarification of the genus Edotea in this regard). Hence, it is apparent from the literature that various workers have interpreted the concept of a ‘‘coxal plate’’ in a variety of ways, some considering a slightly swollen coxa as representing a coxal plate, others assuming lack of an obvious suture line meant the plate had fused with its respective pereonite (when in fact the 268 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON coxa may be so reduced that a dorsal coxal plate simply does not exist). These specific and generic variations are apparently not restricted to the valviferan isopods. Iverson (1978) has recently found that, in the sphaero- matid Exosphaeroma inornata, adult males lack coxal plate sutures entirely, while juvenile males and females bear distinct plates and sutures on per- eonites II through VII. The significant point these discrepancies illustrate is to suggest that the taxonomic value of “‘dorsally visible coxal plates’’ or “‘dorsally visible coxal plate sutures’’ is highly questionable, particularly when relying on the lit- erature. Yet, this character is one that has traditionally been used, often rather casually, in isopod species diagnoses, descriptions, and keys for at least 150 years. Brusca (1977, 1978a, and 1978b) has pointed out that, in the family Cymothoidae, this and similar characters of the coxal plates are of little taxonomic value and often misleading. In the cymothoids the visibility of the plates (in the dorsal view) is often a function of the general body shape, convexity, twisting of the body to the right or left, and state of female gravidness. In our opinion, coxal development can be used as a credible taxonomic character within the Idoteidae only when these structures are considered in their totality: presence or absence of dorsal coxal plates (i.e. dorsolaterally expanded coxae); shape or degree of development; and relative fusion of the coxae with their respective pereonites. We further feel that caution should be exercised in the continued use of the visibility of the coxal plate ‘‘in the dorsal aspect’’ as a reliable taxonomic character, and we urge work- ers to distinguish clearly between ventral and dorsal coxal plates. It is impossible to determine from the literature, for most species, just how many pereonites actually bear coxal plates and/or plate sutures (wheth- er visible in the dorsal aspect or not), and until this particular problem is resolved we do not feel that this character can be considered when eluci- dating a phylogenetic lineage in the higher taxa of idoteid isopods. In the case of Eusymmerus, Parasymmerus, and Colidotea we have been able to examine all but one of the known species (C. edmondsoni) ourselves. We have found that in order to determine adequately whether dorsal coxal plates are actually present, and whether a suture persists, the specimen must often be examined in lateral, ventrolateral, and dorsal aspects, under intense il- lumination (we have had considerable success using a quartz-halogen lamp with a flexible fiber light cord). Key to the Species of Idoteidae Known from the Gulf of California 1. Pleon composed of more than one distinct segment ............... 2 =. Pleon composed of ‘one segment: 4) 6 opie ee Saleen i 2. Pleon composed of 4 segments, plus 1 pair of partial suture lines; flagellum of antenna 2 uniarticulate ........ Cleantioides occidentalis VOLUME 92, NUMBER 2 269 10. Pleon composed of 3 segments, plus | pair of partial suture lines; flaceiiiunnohantenna 2. muttiarticulatey: 402. ask). .eslials. caucuses. 3 + Maxallipedal: palp' ofe4- articles fai4..<4..05%..% Idotea UIdotea) urotoma Meaxllipedalynalprokisvanticles.nisi ont wail. deisk ha setad oes tdeew’h: ie 4 . Eyes transversely (dorsoventrally) elongate and narrow; maxilliped with 1, 2, or 3 coupling hooks ......... Idotea (Pentidotea) stenops Eyes not transversely elongate and narrow; maxilliped with 1 cou- DUNE OUTS Ss OM aes ER Ee eee ge ae a te ere a 5 . Posterior border of pleotelson strongly concave; frontal process extended beyond frontal lamina 1 ...... Idotea (Pentidotea) resecata Posterior border of pleotelson convex, with small median lobe; frontal process not extended beyond margin of frontal laminal .. 6 . Length less than 3.7 times width; eyes reniform; males with distinct tufts of setae on pereopods........ Idotea (Pentidotea) wosnesenskii Length more than 3.7 times width; eyes circular; males without tultis of setac-on pereopeds’ 2 z< 2.2. so. Idotea (Pentidotea) aculeata MrMaceiimeor antenna 2.0f a Single article’... 5.2. P00... ee). 8 Riagetlum ofantenna 2 multiarticulate. ....ic....2.25.5..5026- 5. 10 . Lateral margins of pleon expanded posteriorly; pleon without su- (INKS FIT AS ee eee eee ee Erichsonella cortezi Lateral margins of pleon smooth and gently convex; pleon with one paion partial anterolateral suture lines ©? 222s. 8. ee ed le 9 . Maxillipedal palp of 4 articles; cephalon with a mediodorsal hump; coxal plates of pereonite 6 distinct in dorsal view ............... ee res ee. Aa 8's Mists Oh SOM Ps sas Ge wat CS Eusymmerus antennatus Maxillipedal palp of 3 articles; cephalon without mediodorsal hump; coxal plates of pereonite 6 NOT distinct in dorsal view ... 2a tnagematelemormee! 12 lac cdi ity col Raiden a Parasymmerus annamaryae, N. sp. Maxillipedal palp of 4 articles; posterior margin of pleotelson acu- LITT NT Se ee Oe ee arte Seneca en eee ney See eT eee Colidotea findleyi Maxillipedal palp of 3 articles; posterior margin of pleotelson con- RVI Oe te ie tt NPL elk te garrisoned Bk gies tae Synidotea harfordi Note: The Atlantic species Idotea metallica has not previously been re- ported from the northeastern Pacific; we have, however, a record of a single specimen from the Gulf of California and a second individual, collected by M. Ninos, from Catalina Island, California. This species is commonly found on drifting seaweeds throughout the Atlantic Ocean and elsewhere, and its occurrance in the northeast Pacific is not totally unexpected. ACKNOWLEDGMENTS The authors wish to express their gratitude to Dr. Thomas Bowman and E. Iverson for critically reviewing the manuscript; Phil Pepe and Anna Mary 270 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Mackey for the collection of important materials used in this paper, and other studies; Melinda Thun, for illustrating the dorsal views of S. harfordi, P. annamaryae, and C. occidentalis; Bob Setzer and Dr. David Young, for algal identifications; S. Luke, for assistance in obtaining loan materials from Scripps Institution; and Ruth Toyama, for typing the manuscript. This research was supported, in part, by grants from the National Science Foundation. This is contribution No. 368 of the Allan Hancock Foundation. Literature Cited Benedict, J. E. 1897. A revision of the genus Synidotea.—Proc. Acad. Nat. Sci. Phil. 49:389- 404. Briggs, J. C. 1974. Marine zoogeography.—McGraw-Hill, New York. 475 pp. Brusca, R. C. 1977. Range extensions and new host records of cymothoid isopods (Isopoda: Cymothoidae) in the east Pacific.—Bull. So. Calif. Acad. Sci. 76(2):128—131. . 1978a. Studies on the cymothoid fish symbionts of the eastern Pacific (Isopoda, Cy- mothoidae). I. Biology of Nerocila californica.—Crustaceana 34(2):141—-154. —. 1978b. Studies on the cymothoid fish symbionts of the easter Pacific (Crustacea: Isopoda: Cymothoidae). II. Biology of Lironeca vulgaris.—Occ. Pap. Allan Hancock Found. (n. ser.) 2:1-19. Brusca, R. C., E. Iverson, B. Wallerstein, and B. Winn. (in preparation).—The marine isopods of California. Brusca, R. C. and B. R. Wallerstein. 1977. The marine isopod Crustacea of the Gulf of California. I. Family Idoteidae.—Amer. Mus. Novitates 2634:1-17. Brusca, R. C. and B. R. Wallerstein. 1979. Preliminary comments on zoogeographic patterns of idoteid isopods in the northeast Pacific, with a review of shallow water zoogeography for the region.—Bull. Biol. Soc. Wash. [in press]. Croizat, L., G. Nelson, and D. E. Rosen. 1974. Centers of origin and related concepts.—Syst. Zool. 23(2):265—287. Gurjanova, E. F. 1936. Ravnonogie dalnevostochnykh morei. (Isopoda of far east seas).— Fauna SSSR, Rakoobraznye 7(3): 1-279. Harford, W. G. W. 1877. Description of a new genus and three new species of sessile-eyed Crustacea.—Proc. Calif. Acad. Sci. 7(1):53—55. Iverson, E. 1978. The status of Exosphaeroma inornata Dow and E. media George and Stromberg (Isopoda: Sphaeromatidae) with ecological notes.—Jour. Fish. Res. Bd. Can- ada 35(10): 1381-1384. Johnson, M. E. and H. J. Snook. 1955. Seashore animals of the Pacific coast.—(Reprinted by Dover Publ., New York, 1967) 659 pp. Kensley, B. and H. W. Kaufman. 1978. Cleantioides, a new isopod genus from Baja California and Panama.—Proc. Biol. Soc. Wash. 91(3):658—665. Menzies, R. J. 1950. The taxonomy, ecology and distribution of northern California isopods of the genus /dotea with the description of a new species.—Wasmann Jour. Biol. 8:155— 195. . 1962. The zoogeography, ecology and systematics of the Chilean marine isopods.— Repts. Lund Arsskrift, N.F. 57(2):1-162. Menzies, R. J. and R. J. Waidzunas. 1948. Postembryonic growth changes in the isopod Pentidotea resecata (Stimpson) with remarks on their taxonomic significance.—Biol. Bull. 95:107-113. Menzies, R. J. and D. Frankenberg. 1966. Handbook on the Common Marine Isopod Crus- tacea of Georgia.—Univ. Georgia Press, Athens. 93 pp. VOLUME 92, NUMBER 2 271 Moore, R. C. and L. McCormick. 1969. General features of Crustacea. Jn R. C. Moore (ed.), Treatise on invertebrate paleontology, Part R, Arthropoda 4, pp. R57—R120.—Geol. Soc. Amer. and Univ. Kansas. Miller, M. A. 1975. Isopoda.—/n Smith, R. I. and J. T. Carlton (eds.), Light’s manual. 3rd ed., Univ. Calif. Press. pp. 277-312. Nierstrasz, H. F. 1941. Die Isopoden der Siboga-Expedition. III. Isopoda Genuina. IV. Gna- thiidea, Anthuridea, Valvifera, Asellota, Phreatoicoidea.—Siboga-Expéditie Monogr. 33d:23 1-308. Richardson, H. 1899a. Key to the isopods of the Pacific coast of North America, with de- scriptions of twenty-two new species.—Proc. U.S. Nat. Mus. 21:815—869. 1899b. Key to the isopods of the Pacific coast of North America, with de- scriptions of twenty-two new species.—Ann. Mag. Nat. Hist. 4(7):157—187, 260-277, 321-338. . 1905. Monograph on the isopods of North America.—U.S. Nat. Mus. Bull. 54:1-727. . 1912. Description of a new species of isopod of the genus Cleantis from Japan.— Proc. U.S. Nat. Mus. 42:27-29. Schultz, G. A. 1969. How to know the marine isopod crustaceans.—Wm. C. Brown, Dubuque, Iowa. 359 pp. Sheppard, E. M. 1957. Isopod Crustacea. Part II.—Discovery Rpts. 29:141-198. Tattersall, W. M. 1921. Zoological results of a tour in the Far East.—Mem. Asiatic Soc. Bengal 6:403-433. Wallace, N. A. 1919. The Isopoda of the Bay of Fundy.—Univ. Toronto Studies, Biol. Ser. 18:1-41. Allan Hancock Foundation, Univ. of Southern California, University Park, Los Angeles, California 90007. PROC. BIOL. SOC. WASH. 92(2), 1979, pp. 272-279 A NEW SPECIES OF LEIOCEPHALUS (REPTILIA: IGUANIDAE) FROM HISPANIOLA Albert Schwartz Abstract.—Leiocephalus rhutidira, related to L. vinculum, is described from the southern coast of the northwestern Haitian Presqu’ile du Nord Ouest. Its relationships are apparently with L. v. endomychus of the Haitian Vallée de 1’ Artibonite and Plateau Central. In a series of papers (Schwartz, 1966, 1967, 1968) I reviewed the Hispan- iolan members of the Antillean iguanid lizard genus Leiocephalus Gray. According to Williams’ (1961) concept, the 8 species may be conveniently divided into those that have their centers of distribution on, or are restricted to, the Hispaniolan north island (north of the Cul de Sac-Valle de Neiba plain)—Junatus Cochran, personatus Cochran, pratensis Cochran, schrei- bersi Gravenhorst, and semilineatus Dunn—or on the south island (south of the plain)—barahonensis Schmidt, and melanochlorus Cope. Although L. personatus is widespread on the north island, it also is present in some limited regions on the south island. Geographically, the most peculiar species is L. vinculum Cochran. It is presently regarded as having three subspecies, widely separated from each other: vinculum on Ile de la Go- nave, altavelensis Noble and Hassler on Isla Alto Velo off the southern tip of the Dominican Peninsula de Barahona, and endomychus Schwartz in the interior Haitian Vallée de |’ Artibonite and probably on the Plateau Central. Because of this peculiar distribution, it is difficult to classify L. vinculum as being either north or south island in its affinities. One explanation for the peculiar distribution of L. vinculum is that it was at one time (and perhaps still is, in regions yet unsampled) a widespread lizard which has been forced out of much of its mainland range by more successful competitors, not only Leiocephalus but perhaps also species of the teiid lizard genus Ameiva, and that its present known distribution is a remnant or relictual one resulting from this competition. The often extensive arid habitats on Hispaniola harbor a diversity of moderate-to-large terrestrial lizards whose ecological requirements overlap to a large extent. In the summer of 1978, Eugene D. Graham, Jr., William W. Sommer, and I began an attempt to investigate the herpetofauna of the Haitian Presqu’ile du Nord Ouest, one of the least known regions of Hispaniola. In our attempt we were only partly successful; we were able to travel from Gonaives to Coridon along the base of the southern coast of this peninsula. The road VOLUME 92, NUMBER 2 273 from Gonaives passes over a series of rugged eroded limestone hills, de- scends to the coast at an oasis, and follows the coast through very desolate country at least as far as Coridon. This area is in the rainshadow of the Massif du Nord Ouest, and it is indeed xeric. The rolling hills near Gonaives noted above, although extremely rugged, are somewhat more mesic; this region is in general called Lapierre and is sparsely settled. There are a few scattered small villages or individual houses along the roadway, and it was relatively simple to encourage people to collect herpetological specimens for us. We have little expectation of anything exciting since the area is bleak, but we were amazed at the diversity of the herpetofauna there. The dominant Leiocephalus, seen sunning on rocks and running across the road, is L. schreibersi; this is certainly the expected member of the genus in such a habitat at this locality. On 9 July a single adult male of a very different style of Leiocephalus was brought to us. We urged the natives to secure more of these, and they obtained five additional specimens. They indicated that the lizards occurred at the base of the hills above the village. We never saw any of them ourselves, and the precise habitat occupied by these lizards remains conjectural. The new lizards are closely allied to L. vinculum and might properly be regarded as a subspecies of that species, but the hiatus in the ranges and the characteristics of the new lizards suggest that they are a L. vinculum derivative, probably limited to special ecological situations on the Presqu ‘ile du Nord Ouest—in other words, another fragment of a basic and now much splintered L. vinculum stock. Until much more is known of the variation and especially the distribution of L. vinculum on Hispaniola itself, specific status for these lizards is the more appropriate alternative. Leiocephalus rhutidira, new species Fig. 1 Holotype.—Carnegie Museum of Natural History (CM) 60520, adult male, from Lapierre, 10.6 km W Ca Soleil, 122 m, Département de I’ Artibonite, Haiti, one of a series collected by native collectors, 14 July 1978. Original number Albert Schwartz Fieid Series (ASFS) V46747. Paratypes.—ASFS V46748-50, same data as holotype; ASFS V46324, same locality as holotype, 9 July 1978; ASFS V46742, 6.7 km W Ca Soleil, 61 m, 14 July 1978. Definition.—A species of Leiocephalus characterized by a combination of: 1) small size (males to 66 mm, females to 56 mm snout-vent lengths); 2) distinct sexual dichromatism, and males with a prominent pair of enlarged postanal scales; 3) absence of a lateral fold; 4) dorsal scales imbricate, not denticulate or tricuspid, and keeled; ventral scales imbricate, smooth, and not denticulate; 5) median crest scales slightly enlarged, lower than median 274 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Qe ee Rebiae Onspe/2 _~. rF eheeeelde ol ARES MATS Raa ep sate yo EE ty Fig. 1. Lateral and ventral views of the holotype of Leiocephalus rhutidira (CM 60520). dorsal caudal scales, 51—65 in occiput to vent distance; 6) one-half midbody scales 22—25; 7) supraoculars 5/5 to 6/6 (no mode); 8) loreals 4—6; 9) tem- porals 8-10; 10) supraorbital semicircles usually incomplete; 11) parietals always in contact; 12) median head scales either 4 or 5, mode 4; 13) pre- auricular scale small (see Schwartz, 1967:4 for this usage); 14) throat, chest, anterior faces of forelimbs, and sides of neck between prominent neck wrin- kles black in males; 15) ventral color orange in males, pale yellow in females; 16) ventral pattern absent in both sexes except for scattered greenish scales VOLUME 92, NUMBER 2 ZS on the sides of the abdomen in males; 17) facial mask absent in males, poorly shown in females where it is the anteriormost pale portion of a broad lateral dark gray band which extends from the hindlimbs to the eyes; 18) dorsum unpatterned tan to grayish tan in males with a conspicuous pair of broad pale cream to pale grayish tan dorsolateral stripes from the supraor- bital semicircles posteriorly along the length of the body, becoming paler posteriorly; 19) throat in females and juveniles with a gray reticulum with vague indications in this pattern of a pair of paramedian gray lines. Description of holotype.—The holotype is an adult male with a snout-vent length of 66 mm and a broken tail; the scale counts are: dorsals occiput-vent 59, dorsals occiput-axilla 24, trunk dorsals 35; one-half midbody scales 24; tricarinate subdigital scales on fourth toe 24 on right foot, left foot uncount- able; loreal scales 5/6, supraocular scales 5/5, temporals 10, supraorbital semicircles incomplete, parietals in contact, 3 prefrontal scales, 4 median head scales, 5 frontoparietal scales, the prefrontal and frontoparietal rows complete. Dorsum in life dull grayish tan and without chevronate pattern; a pair of faintly paler longitudinal lines, 3 to 4 scales wide, from behind the eyes posteriorly to above the hindlimb insertions; all dorsal head scales tan with more or less symmetrical black spotting on the paraparietals and the parietal, the supraorbitals, and the snout, those on the snout mere flecks; uppersides of hindlimbs pale green, those of forelimbs concolor with dor- sum; throat, chest, and anterior faces of forelimbs black, with a very few scattered greenish scales on the chin and throat and more abundantly at the junction of the black chest area and the orange venter; scattered iridescent bright green scales on the lower sides, which are pale grayish tan, and laterally onto the venter; black throat pigment extending dorsally between the conspicuous neck wrinkles to form three extensions of this color on the sides of the neck, the wrinkles themselves set off from the balance of the neck color by white scales, no dark mask present but a subocular black dot, intensified by a white dot on each side, as well as a white dot at the posterior corner of the eye. Variation.—Scale counts for the series (including the adult male holotype, a subadult male, an adult female, a subadult female, and two juveniles) are: dorsal scales occiput-vent 51-65 (*& = 58.4), dorsal scales occiput-axilla 19-— 24 (20.8), dorsal trunk scales 32—44 (37.8); one-half midbody scales 22—25 (23.7); fourth toe subdigital scales 18—23 (21.7); loreals 4—6 (5.0); temporals 8—10 (9.0); supraocular scales 5/5 (2 individuals), 5/6 (2), 6/6 (2); supraorbital semicircles incomplete (4 individuals) or complete (2); prefrontal scales 2 or 3 (M, = 3, 66%), row always complete; median head scales 4 or 5 (M, = 4, 83%); frontoparietal scales 3—5 (M, = 5, 50%); modal head scale formula 3— 4_5, The subadult male (ASFS V46324; snout-vent length 59 mm) was de- scribed in life as ‘‘dorsal ground color tan, with a conspicuous pair of pale 276 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON cream dorsolateral stripes on neck and anterior part of trunk; head shields bright tan with symmetrical black flecking on paraparietals and supraocu- lars; neck and shoulders and supraaxillary areas with three jet black patches. Upper sides of all limbs concolor with tan of dorsum, flecked with pale tan; throat deep orange, all other ventral surfaces yellow including underside of tail; upperside of tail tan, dorsal crest scales cream; tail banded tan-and- cream.’’ After preservation, the throat is dark dusky gray and the black interwrinkle dorsad patches noted above arise from this dark throat area. The dorsolateral stripes are more prominent in this subadult male than in the holotype. The differences in coloration and intensity between the sub- adult and adult males are probably related to the age difference. The adult female (ASFS V46748; snout-vent length 56 mm) was described in life as: ‘‘dorsal ground color dark brown with a pair of buffy dorsolateral stripes; sides very dark gray; ventral ground color pale yellow; pale grayish orange on throat, with a vague dark gray reticulum and indications of a pair of paramedian longitudinal gray lines.’’ The neck wrinkling is apparent on the female but there are no black patches between the wrinkles nor are they set off by white coloration. Aside from the dorsolateral stripes, this female is a very dull colored and drab lizard. The dark gray sides, bordered above by the buffy dorsolateral stripes and below by a vague paler longitudinal stripe between the fore- and hindlimbs, are especially prominent. The subadult female (ASFS V46742; snout-vent length 45 mm) was like the adult female except that the dorsum was tan and the venter yellow with a more orange tinge on the throat. The throat pattern was a pale gray retic- ulum, and the paramedian gray lines are so broken as to be hardly recog- nizable. ; The two juveniles (ASFS V46749_-S0) have snout-vent lengths of 30 mm. They are colored and patterned like the females except that the dorsolateral Stripes are white anteriorly, strongly contrasting with the tan to brown dor- sal color, and the ventral ground color is yellow-green. The sides are very dark gray and accentuate the white dorsolateral stripes. The throat markings are a dark gray reticulum, but the throat ground color is not orange. One juvenile (ASFS V46750) has indications of about five faint, widely spaced, darker brownish transverse bands across the back between the longitudinal stripes and is the only specimen with any sort of middorsal pattern. Comparisons.—The details of scutellation and of color and pattern elim- inate all other Hispaniolan (or even Antillean) Leiocephalus as near rela- tives of L. rhutidira with the exception of L. vinculum. Of the Hispaniolan species, L. melanochlorus and L. schreibersi have lateral folds, L. pratensis lacks a median dorsal row of crest scales, L. barahonensis and L. semilin- eatus have enlarged preauricular scales, L. /unatus males have white throats with distinct black dots, and L. personatus females have grayish throats with darker gray dots. All these species also differ in many other ways from VOLUME 92, NUMBER 2 277 L. rhutidira as well. The key to the Hispaniolan species of Leiocephalus in Schwartz (1967:52—53) reaffirms these distinctions. That L. rhutidira is clos- est to L. vinculum is obvious in that, when specimens of the former are run through the key, they will key out to L. vinculum, provided one uses the key with some latitude. One key-character for L. vinculum is the presence in both sexes of a dark throat (heavily patterned and on a dusky ground in some subspecies), whereas female L. rhutidira have a dusky throat with a vestigial or faint pattern which is much like the juvenile and subadult female patterns in some subspecies of L. vinculum. One pattern feature of L. rhutidira is the lack of any sort of dorsal cross- banding, marbling, dotting, or other sorts of markings found in L. vinculum. Leiocephalus rhutidira is a less ornate lizard dorsally than is L. vinculum. The solid dark gray sides of female L. rhutidira differ markedly from those of L. vinculum females. Leiocephalus rhutidira shares with L. v. vinculum the longitudinal paired stripes, but in the latter the throat is not solid black nor are there lateronuchal black extensions. Leiocephalus v. endomychus, the only mainland subspecies of L. vincu- lum, was described on the basis of a male and a female from near the Barrage de Peligre in the Vallée de l’Artibonite. Another specimen from Hinche on the Plateau Central was tentatively associated with the Peligre material. Since that time, a series of 15 more near-topotypes of L. v. en- domychus has been secured, so the subspecies is now well represented in collections, and its variation can better be assessed. Field notes in life for a male (ASFS V43786; snout-vent length 66 mm, the same size as the ho- lotype of L. rhutidira) state that the ‘‘dorsal ground color is green with gray chevrons, the limbs not brighter than the dorsum; the head was brownish, the throat and chest jet black, with some scattered pale green dots; the ventral ground color was greenish orange.’’ Another male (ASFS V43787; snout-vent length 69 mm) was recorded as being like the male described above but with the dorsal chevrons somewhat less obvious. Neither speci- men (nor the male holotype) has dorsolateral stripes. Females are tan to gray above, with limbs slightly greenish; the ventral ground color is pea green and the gray throat is heavily spotted and has a central pair of lon- gitudinal lines. As far as can now be determined, neither sex has the strongly wrinkled neck of L. rhutidira, although the black throat and chest color of males does extend dorsally onto the sides of the nuchal area as in L. rhu- tidira. Etheridge (1966:85) pointed out that nuchal and antebrachial folds are a diagnostic character of Leiocephalus, but their development is some- what variable within the genus. Even the smallest L. v. endomychus (ASFS V43794; snout-vent length 35 mm) lacks the dorsolateral stripes so conspic- uous in juvenile L. rhutidira. Considering all three subspecies of L. vinculum together, maximum snout-vent length of males varies between 78 mm (vinculum) and 69 mm 278 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON (endomychus), and that of females ranges between 73 mm (vinculum) and 63 mm (altavelensis). The largest L. rhutidira are smaller than any of the subspecies of L. vinculum; this may of course be a sample artifact, but the largest male L. rhutidira (66 mm) already has the distinctive coloration and pattern. Dorsal crest scales occiput to vent in L. vinculum vary between 51 and 65, with means of 55.8, 56.7, and 57.2; the range in L. rhutidira is 51- 65, with a mean of 58.4, slightly greater than that of any subspecies of L. vinculum. The same is true of dorsal crest scales in occiput-axilla, with a mean of 20.8 in L. rhutidira and means of 18.6 to 20.0 in the subspecies of L. vinculum. Trunk scales show the same relationship, although the mean for L. rhutidira (37.8) is only slightly greater than that for L. v. altavelensis (37.7). The mean of 5.0 loreal scales in L. rhutidira is greater than the means for the subspecies of L. vinculum (3.7-—4.8). In L. vinculum, the supraorbital scales vary between 4/5 and 7/8, with ‘‘abnormal’’ counts in two specimens of 6/8 and 9/10. Modes in all cases are 6/6 (47% in L. v. endomychus to 96% in L. v. vinculum). But although the modes are the same in all subspecies (and in fact 6/6 in the “‘normal’’ count in most species of Leiocephalus), L. v. endomychus differs from L. v. vinculum and L. v. altavelensis in having low counts (4/5—S/6) in 9 of 17 specimens, whereas only one specimen of a total of 77 L. v. vinculum and L. vy. altavelensis has a count less than 6/6. Leiocephalus rhutidira has counts resembling those of L. v. endomychus—S/5, 5/6, 6/6, all with equal frequency. In supraorbital semicircles, 4 of 6 L. rhutidira (67%) have these series incomplete, whereas they are complete in 69% of L. v. vinculum and 94% of L. v. endomychus; the series of L. v. altavelensis is equally divided. The head scale formula of L. v. vinculum, L. v. altavelensis, and L. rhutidira is 3-4-5, but L. v. endomychus has the formula 2—4—4. Prefrontal rows are complete in both species; frontoparietal rows are usually complete in all taxa (always so in L. v. altavelensis and L. rhutidira) but are incomplete in 23% of L. v. vinculum and 6% of L. v. endomychus. The usual number of prefrontals in Leiocephalus is 3, but only 2 are present as a variant in many species, with low frequency or as an aberrant condition. This is not known in L. v. altavelensis, but 36% of L. v. vinculum, 59% of L. v. endomychus, and 67% of L. rhutidira have but 2 prefrontals. Etymology.—The name rhutidira is from the Green rhutis (wrinkle) and deire (neck), in allusion to the prominent neck wrinkles, and is a noun in apposition. Remarks.—QCa Soleil is unlocatable on many maps. It is the local name for the intersection of the major north-south road between Gonaives and Cap-Haitien, with the coastal road to the northwest toward Coridon and the Presqu ile du Nord Ouest. Specimens examined.—All specimens of L. v. vinculum and L. v. alta- VOLUME 92, NUMBER 2 279 velensis whose data have been used in the present paper are listed in Schwartz (1967). Additional specimens of L. v. endomychus utilized in the present study are from: Haiti, Dépt. de l’Artibonite, 1.1 km—5.1 km NE Barrage de Peligre, 180 m—335 m (ASFS V43786, V43787-93, V43794, V43795-96, V43800, V43801-03). Acknowledgments The series of L. rhutira is the result of industry on the part of Haitians at Lapierre, but my companions Eugene D. Graham, Jr., and William W. Sommer made the visits to that region pleasant and profitable. The series of L. v. endomychus was collected in 1976 through the efforts of David A. Daniels and William W. Sommer. The illustrations are the work of Beryl Bayer. Literature Cited Etheridge, R. 1966. The systematic relationships of West Indian and South American lizards referred to the iguanid genus Leiocephalus .—Copeia (1):79-91. Schwartz, A. 1966. The. Leiocephalus (Lacertilia, Iguanidae) of Hispaniola. I. Leiocephalus melanochlorus Cope.—J. Ohio Herpetological Soc. 5(2):39-48. . 1967. The Leiocephalus (Lacertilia, Iguanidae) of Hispaniola. II. The Leiocephalus personatus complex.—Tulane Stud. Zool. 14(1):1-53. 1968. The Leiocephalus (Lacertilia, Iguanidae) of Hispaniola. III. Leiocephalus schreibersi, L. semilineatus, and L. pratensis.—J. Herpetology 1(1—4):39-63. Williams, E. 1961. Notes on Hispaniolan herpetology. 3. The evolution and relationships of the Anolis semilineatus group.—Breviora 136:1-8. Miami-Dade Community College North, Miami, FL 33167. PROC. BIOL. SOC. WASH. 92(2), 1979, pp. 280-286 A NEW SPECIES OF TREE SQUIRREL (SUNDASCIURUS) FROM PALAWAN ISLAND, PHILIPPINES (MAMMALIA: SCIURIDAE) Lawrence R. Heaney Abstract.—A new tree squirrel, Sundasciurus rabori, new species, from the mountains of Palawan Island, Republic of the Philippines, is named and described. It is assigned to the subgenus Aletesciurus on the basis of cranial characters and morphometrics. Relationships of taxa within the genus are considered briefly. In 1962, an expedition sponsored by the U.S. Naval Medical Research Unit No. 2, Bernice P. Bishop Museum of Honolulu, and the Silliman Uni- versity of Dumaguete City, Republic of the Philippines, collected verte- brates on Palawan and nearby smaller islands in the southwestern portion of the Philippines (Kuntz, 1969). Among the 863 mammals obtained, all of which are now deposited at the National Museum of Natural History, Smith- sonian Institution (USNM), are five squirrels representing an undescribed species that differs markedly from all other Philippine tree squirrels. This paper describes this new species and briefly documents its relationship to other members of the genus Sundasciurus. Methods Cranial measurements were made with dial calipers graduated to one- twentieth of a millimeter (mm). All capitalized color names are from Ridg- way (1912). All measurements are as defined in DeBlase and Martin (1974) except the following: rostral depth, taken from the point where the maxil- lary-premaxillary suture crosses the midline on the ventral surface of the rostrum, to the closest point on the midline of the dorsum of the rostrum; rostral length, taken from the anteriormost point of the nasals at the midline to the closest point in the orbit; orbital length, taken from the point in the orbit where the preceding measurement was taken (i.e., the anteriormost) to the most posterior point in the orbit (see Moore, 1959:164); palatal breadth, taken from the labial edges of P4; length of diastema, taken along the midline from the posterior margin of the incisors to the line defined by the anterior points of the maxillary toothrow. External measurements shown in Table 1 were taken from specimen labels. For S. hippurus and S. brookei the total length shown is the sum of head and body length plus tail length, VOLUME 92, NUMBER 2 281 as recorded by the collector. A cluster analysis (BMDP2M) and principal components analysis (BMDP4M) were performed on a Honeywell 66-60 computer at the University of Kansas. Comparisons of taxa are based on specimens at the USNM, but data used in compiling Table 1 were also obtained from specimens housed in the Field Museum of Natural History (FMNH), as noted in the text. Sundasciurus rabori, new species Holotype.—Adult male represented by skin and skull, USNM 477989. Obtained 16 April 1962 at Magtaguimbong, Mt. Mantalingajan, Palawan Is- land, Republic of the Philippines, between 3,600 and 4,350 ft elevation by D. S. Rabor; this is approximately 8°48’N, 117°40’E. Skull complete and unbroken; skin well prepared, with terminal portion of tail missing, though the external measurements taken by the collector were probably taken prior to this loss. Specimens examined.—Skins and skulls of five individuals, all from the type locality, USNM 477985-—477989. Numbers 477985 and 477989 are adults; the skull of 477985 is crushed. Numbers 477986—477988 are in adult pelage but had not completed growth, as indicated by thin cranial bone and open basioccipital sutures; I consider these subadults. Measurements.—External measurements of the holotype, the second adult male, and the mean for the three subadults, all taken from the labels made out by the collector, and my cranial measurements are listed in Table 1. Diagnosis.—Dorsal and lateral fur dense and soft, mostly 15 mm long, colored a dark brown agouti. Terminal bands black, subterminal bands cor- responding most closely to Ochraceous Orange, and basal bands black. Subterminal bands of hair on nose paler than elsewhere, corresponding to Pale Yellow-Orange. Color of the hair on the ears and around the eyes not different from that of the surrounding hair. Ventral region, especially within 1 cm of the midline from the throat to the anal region, and ventral surface of the limbs washed with silver, corresponding to Light Buff. Tail fur agouti with three alternating bands of Ochraceous Orange and black, and tipped with Pale Yellow-Orange. Skull (Fig. 1) relatively gracile; rostrum slender; postorbital region short; temporal ridges not fused to form a sagittal crest; posterior portion of the cranium domed, not curving evenly to the occipital crest; palate narrow, strongly concave. Bullae small, with a single septum which is divided to form a small anteromesial lobe, with no portion con- spicuously inflated. Additional specimens examined.—S. h. hippurus (6), Gunong Tebu (350 ft), Trengganu, West Malaysia, USNM 311342-311345, Bukit Besi, Treng- ganu, West Malaysia, USNM 311346-311347; S. hoogstraali (10), 6 km NE San Nicolas, Busuanga Island, USNM 477850-—477857, 477860, 477862; S. 282 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 1. Ventral and dorsal views of the skull of the holotype (USNM 477989) of Sundasci- urus rabori, approximately twice life-size. jJuvencus (23), Tarabanan, Concepcion, Puerto Princesa, Palawan Island, USNM 477879-477883, 477890—477898, 477902—477910; S. mindanensis (2) Loreto, Surigao del Norte, Dinagat Island, USNM 462199—462200; S. mol- lendorffi (2), 6.5 km SW Culion, Culion Island, USNM 477946-477947; S. samarensis (16), Samar or Leyte Island, USNM 105464—105465; Matugui- nao, Samar Island, FMNH 87722-87726, Mt. Capoto-an, Samar Island, FMNH 87728-87736; S. steerii (21), Minagas Point, Dalawan Bay, Balabac Island, USNM 477964—477984; S. brookei (2), Poring, Ranau National Park, Sabah, USNM 488399—488400; S. 7. jentincki (7), Lumu Lumu, Mt. Kina- balu, Sabah, USNM 292570-292576; S. 1. lowii (5), Sungei Djambajan, Bor- neo, USNM 198739, 198744—198745, Ranau, Sabah, USNM 300988-300989, S. t. tenuis (10), 16 km N Kuala Lumpur on Pahang Road, Selangor, West Malaysia, USNM 283482-—283483, 290160—290167. Comparisons.—Moore (1958) named the genus Sundasciurus on the basis of characters of the bulla, noting that his classification showed some cor- respondence with that of Thomas (1915) based on the structure of the bac- 283 VOLUME 92, NUMBER 2 yO= TOF £0 70+ LO €£0F V0+ LOF £0 SOF 90+ 90+ 96F STF OCF TL+ BCI+ snaniosppung 98 O8 79 ITU OST DL TS TI €9l 9% LT Mee O€B HET Tle ETIl OOSe ZL sinua} “1 *S £O+ CO} £0 [0 60+ COy F090] 70-90 V0 ls So eee ae Lee SHAMIISOPUNS SCG) O'S 19s Sch 6 S12 98 SCS SIE eGiCCee Ue SS ann OLS TS ayo 8528) = ORBCCe Ss Gah 90+ VO+ TOF £0+ SOF TOF £0+ VO POF SOF 10 80F elas LeaiO Oke snanjosopunS 78 vL O99 BIl Svl 69 6h 6OL BST DIZ VU tte — OI CHE BOIL LHe S ryounual *f snanosppuns 96 68 OL SEI PLE $8 8S LU E81 Tse Ll OLE El bl 9 CEL 862 ¢ ROO ARES: pO+ L0+ 0+ VO+ SO €0* TOF S0+ 90+ 60+ 80F 60F | Bat Sac (Gear A ae snamgsaiaqy V7l TIL 66 LSt 7% VOI SL 9 OC HOE ELI DLb — Y6I 6&h HIST 6L9E El Mdaa}S. “S 90+ 20+ £0F CO= 90+ F000 90 © 0-0 S093 05 SO ORC CS ay snaniosajaqqv CL SOL 98 EST 907 COI 89 SSt C17 967 OLI Sh — eC8L T9r LHOl Bese cl SISUQAD IDS. 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Cluster phenogram of morphometric similarity of 12 species of Sundasciurus, based on data in Table 1. ulum. He divided the genus into two subgenera. The subgenus Aletesciurus included all named Philippine forms, as well as S. hippurus from the Sundaic Subregion. Diagnostic features are their larger size (skull length over 50 mm according to the original description, but over 45 mm according to Moore, 1959), presence of an inconspicuous anteromesial lobe in the bulla, and presence of a sagittal crest in adults. The nominate subgenus contained four species, all from the Sundaic Subregion, which have skull lengths under 40 mm, an inflated anteromesial lobe, and no sagittal crest. Sundasciurus rabori is intermediate in size between the two subgenera as defined by Moore (1958), although closer to Aletesciurus (Table 1). In com- mon with the small species, S. rabori lacks a sagittal crest; in most sciurids I have examined there is a positive allometric relationship between size and development of the cranial crest, and I hesitate to use the crest as an indi- cator of reiationship without further study. Each bulla of S. rabori has a small anteromesial lobe which is inflated more than in S. hippurus or the Philippine forms, but is conspicuously less inflated and less sharply defined than in the smaller Sundasciurus. Moore (1959) found that the structure of the bulla was a consistent and useful character for discerning relationships among diurnal squirrels. I conclude that, although intermediate in some ways, S. rabori is more similar to the subgenus Aletesciurus than Sunda- sciurus on the basis of the discrete characters available. To investigate the relationships of S. rabori further, I studied the phenetic VOLUME 92, NUMBER 2 285 similarity of 12 taxa using a cluster analysis based on the cranial measure- ments in Table 1, using only the type of S. rabori for this analysis. The resulting phenogram (Fig. 2) supports the division of Sundasciurus into two subgenera as defined by Moore (1958) and discussed above. Within the subgenus Aletesciurus the greatest difference is between S. hippurus and the Philippine members of the subgenus. S. rabori is most similar to the lowland Philippine squirrels, which are themselves divisible into a southern Philippine group (containing S. mindanensis and samarensis, and probably S. philippinensis and S. davensis—not examined), and a Palawan group (containing S. hoogstraali, S. juvencus, S. mollendorffi, and S. steerii). It is possible that the southern Philippine group and the Palawan group each represent a species with several subspecies, but further analysis is required before this taxonomic change can be justified, and listing of S. juvencus as a subspecies of S. steerii by Kuntz (1969) is premature. I conclude from the above analyses that S. rabori is most closely related to the other Sundasciurus from the Philippines, and should be assigned to the subgenus Aletesciurus. A principal components analysis identified length of the diastema, orbital length, rostral depth, length of the maxillary molariform toothrow, and pal- atal breadth as the five measurements most useful for distinguishing among all taxa. S. rabori was characterized by this analysis as having a relatively short toothrow, narrow palate, long diastema and orbit, and deep rvstrum. Remarks.—Although specimens of S. rabori have been taken only on Mt. Mantalingajan, which is the highest mountain on Palawan at 6,841 ft (2,085 m), they may occur widely on Palawan at elevations above 1,100 m, which is the elevation at which a major vegetational change takes place in most of Southeast Asia (Medway, 1972; Steenis, 1964). Regions above this elevation include substantial areas in the Mantalingajan Range, Victoria Range, and Cleopatra Needle. No other island in the Palawan chain has mountains over 765 m. No information is available regarding the ecvlogy of this species. Etymology.—This species is named for its collector, Dr. Dioscoro S. Ra- bor, in recognition of his contributions to knowledge of the vertebrates of the Philippines. I suggest ‘‘Palawan montane squirrel’ as an English name. Acknowledgments This paper benefited from suggestions and editorial comments from Greg- ory Glass, Robert Hoffmann, James Koeppl, Thomas McIntyre, Gary Mor- gan, Guy Musser, Richard Thorington, and Robert Timm. Partial support was provided by the Smithsonian Institution Office of Awards and Grants. Computer facilities were provided by the Academic Computer Center of the University of Kansas. Richard Thorington and Patricia Freeman provided access to specimens housed in the USNM and FMNH, respectively. 286 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Literature Cited DeBlase, A. F., and R. E. Martin. 1974. A manual of mammalogy.—W. C. Brown Publ. Co., Dubuque. xv + 329 pp. Kuntz, R. E. 1969. Vertebrates taken for parasitological studies by U.S. Naval Medical Re- search Unit No. 2 on Silliman University—Bishop Museum Expedition to Palawan, Re- public of the Philippines.—Quart. J. Taiwan Mus. 22:207-220. Medway, Lord. 1972. The Quaternary mammals of Malesia: a review. pp. 63-83 in P. Ashton and M. Ashton (eds.), Transactions of the Second Aberdeen—Hull Symposium on Male- sian Ecology.—Univ. Hull, Dept. Geography, Misc. Series no. 13. 122 pp. Moore, J. C. 1958. New genera of East Indian squirrels —Amer. Mus. Novitates 1914:1-S. . 1959. Relationships among the living squirrels of the Sciurinae.—Bull. Amer. Mus. Nat. Hist. 118:153—206. Ridgway, R. 1912. Color standards and color nomenclature. Washington, D.C.—Published by the author. ii + 43 pp, 53 pls. Steenis, C. G. G. J. Van. 1964. Plant geography of the mountain flora of Mt. Kinabalu.— Proc. Roy. Soc. London, Series B. 161:7-38. Thomas, O. 1915. The penis-bone, or ‘‘baculum,”’ as a guide to the classification of certain squirrels.—Ann. Mag. Nat. Hist. (ser. 8) 15:383-—387. Museum of Natural History and Department of Systematics ang Ecology, University of Kansas, Lawrence, Kansas 66045. PROC. BIOL. SOC. WASH. 92(2), 1979, pp. 287-293 NEW SPECIES OF NEARCTIC HYDROPORUS (COLEOPTERA: DYTISCIDAE) James F. Matta and G. William Wolfe Abstract.—Two new species Hydroporus sulphurius and Hydroporus ouachitus are described from the interior Highlands of Arkansas and a third new species, Hydroporus alleghenianus, is described from the Applachian Highlands of Virginia and Tennessee. All three species are placed in the pulcher group of Hydroporus. Introduction The pulcher group of the genus Hydroporus was first keyed and the species discussed by Fall (1923) in a revision of the entire genus Hydropo- rus. Since that revisionary work one new species has been described in the pulcher group by Leech (1949) and three species by Wolfe and Matta (1978). While going through the material at the National Museum of Natural History the authors discovered an additional new species in this group from Sulphur Springs, Arkansas. During a collecting trip to Arkansas to obtain additional material of this species a second new species in this group was collected. These two species and a third species which has been in the senior author’s collection for several years are described below. Hydroporus sulphurius, new species Diagnosis.—This species will key to the first part of couplet 7 in Fall (1923) but does not fit either of the species listed there. The completely infuscate pronotum separates it from all species which will key to this cou- plet except H. laetus, which is similar in size, pronotal markings and lack of prosternal setae. The punctation of the two species is similar but the punc- tures of H. laetus are slightly larger and more densely applied than that of AZ. sulphurius; in addition the elytral markings of sulphurius are darker than those of /aetus. The aedeagus is distinct and may be used to separate this species from all others in the group (see Fig. la, b). Description.—Holotype: Male: length 3.15 mm, width 1.55 mm, L/W = 2.03. Form elongate oval. Lateral margins of the pronotum evenly rounded towards the anterior angles, with a distinct bead which appears flattened on top and which is about one half as wide as the last antennal segment. Pros- ternal setae absent; prosternal process with an angulate protuberance and a poorly defined anterior file. Posterior to the procoxae the prosternal pro- 288 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON a b Cc d @ f Fig. 1. A. sulphurius, aedeagus, a—dorsal, b—lateral; H. ouachitus; aedeagus, c—dorsal, d—lateral; H. alleghenianus, aedeagus, e—dorsal, f—ateral. cess is lanceolate shaped, rounded medially, with a well developed flattened lateral margin and is produced to a bluntly rounded tip. ; Head pale brown; antennae concolorous with head, vaguely infuscate on last segment. Pronotum rather uniformly reddish brown; lateral margins lighter. Each elytron reddish brown with three light brown pale patches which extend across the width of the elytron (except for a narrow dark band at the suture); one at the anterior margin, one posteriomedially and one at the apex. The posteriomedial spot is interrupted by a narrow dark band cutting across its middle and the apical spot extends anteriorly as a vague narrow streak but does not reach the posteriomedial spot. Ventral surface light brown with the metacoxal plates, metasternum and prosternal process slightly darker. Head sparsely punctured, the punctures quite fine. Pronotum with a nar- row band of coarse punctures near the anterior margin and a somewhat broader band of coarse punctures on the posterior margin. The disc is as finely punctured as the head. Elytral punctures fine and sparsely distributed, Separated by at least the width of one puncture, usually by the width of several punctures. The entire dorsal surface is finely and distinctly aluta- ceous between the punctures; the individual spaces outlined by the reticu- lation about the size of a puncture. Ventral surface finely punctured except for the metacoxal plates and metasternum where the punctures are 4—5 times larger than normal. The protarsal claws are unmodified and the protarsi do not exhibit any appreciable sexual dimorphism. The aedeagus is apically bifid and 1s illus- trated in figure la, b. Allotype.—Length 3.1 mm, width 1.55 mm, L/W = 2.00. The dark mark- ings of the elytra and pronotum are slightly darker than those of the male. VOLUME 92, NUMBER 2 289 The allotype is otherwise similar to the male in color, punctuation and body form. Variation.—Since the sexes are not easily distinguished on the basis of external morphology the variation in the type series is not reported by sex. Average length 3.24 mm (2.9-3.5); average width 1.55 mm (1.4-1.7); L/W = 2.087; n = 20. The color pattern is remarkably constant within the type series. The pronotum is usually concolorous; however a few specimens have a vague paler area on the disc. Some specimens have a dark spot at the anterior edge of the anterior elytral pale spot, some have the post median pale spot reduced or split by a dark streak. The dark areas on the ventral surface vary in extent in the type series from the typical reduced dark areas to large infuscate patches on the ventral abdominal segments. Habitat.—The type locality is located on the Ozark plateau in an area of numerous springs, and small sand bottom streams. No specific habitat data is available for this species; however it is probably found at the margins of the smaller streams. The collection from which the type series was obtained contained a large series (280) of Hydroporus wickhami Zaitzev. Type data.—The holotype, allotype and 31 paratypes are from Sulphur Springs, Arkansas; VII-20-1955; P. J. Spangler. Etymology.—The authors wish to thank Dr. Paul Spangler for suggesting the name for this species. It is named after the type locality and also for the mood of the senior author when he failed to collect additional specimens during a visit to the type locality in the summer of 1978. Deposition of type material.—The holotype and allotype are deposited at the National Museum of Natural History (NMNH) and are assigned type number 97565. Sixteen paratypes have also been deposited in the NMNH, two paratypes with the Canadian National Collection (CNC) and 13 para- types have been retained by the authors (6 by JFM and 7 by GWW). Hydroporus ouachitus, new species Diagnosis.—This species is similar to H. pulcher and H. cocheconis and will run to those species in existing keys. It may be separated from those species by the metasternal and metacoxal punctation which is coarser than pulcher but finer than cocheconis; by the smaller size, the reduction of the anterior pronotal dark spot and by the shape of the male genitalia. In ad- dition all of the type material has the last antennal segment partially infus- cate. This is a variable character in pulcher but the last antennal segment is usually not infuscate and was not infuscate on any specimens which are collected with the type series. Description.—Holotype: Male: Length 2.82 mm, width 1.3 mm, L/W ratio 2.7. Form elongate oval, widest at the middle; with the sides of the elytra very slightly concave basally. Lateral margin of the pronotum with a distinct 290 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON lateral bead which tapers slightly in the posterior fourth. Lateral edge of the pronotum evenly rounded towards the anterior angles. No prosternal setae present although a few short stiff fine hairs are present on the prosternum at the base of the coxae. Prosternal process distinctly angulate and protu- berant, the prosternal file present but very poorly developed. The posterior half of the prosternal process broadly lanceolate with the medially portion slightly rounded and lateral edges margined; the tip bluntly rounded. Head reddish brown, antennae with the last segment infuscate. Pronotum reddish brown, with a diffuse dark spot at the anterior margin which reaches from the central line to approximately half the distance to the margin on each side. Posterior margin of pronotum with a thin dark band which nar- rows laterally. Elytra dark brown with subbasal, postmedial and apical pale areas which are separated from the suture and the lateral margin by thin dark bands. Ventral surface yellow-brown, the legs, prosternum and head reddish brown. Dorsal surface finely and densely microreticulate; punctation of head fine, separated by 3-4 times the width of the punctures. Pronotum with discal punctures similar to those of the head; with a band of coarser punctures near the anterior margin and with a few coarse punctures scattered along the posterior margin. Elytra rather evenly punctured, the punctures sepa- rated by 1-1'% puncture diameters. The elytral punctation is intermediate to H. pulcher and H. cocheconis. Ventrally metacoxae and metasternum coarsely punctured, the punctures separated by less than 2 a puncture diameter. The abdominal punctures fine, slightly coarser basally. Pro- and mesotarsi short and thickened, the first three segments with a glandular pubescence and small palettes. The anterior protarsal claw is not modified. Aedeagus bifid and is in figure 1b. Allotype.—Length 3.0 mm, width 1.45 mm, L/W ratio 2.07. Similar to the male except as follows: slightly longer and broader, the anterior dark spot of the pronotum is less diffuse; the dark markings of the elytra are reduced, the subbasal pale spot not separated from the margin by a dark band. Last segment of antennae infuscate only at the tip. Ventral punctation slightly finer. Variation.—Average length 2.94 mm (2.80-3.20); Average width 1.40 mm (1.35-1.45); L/W = 2.10; (n = 10). Males average 0.1 mm shorter than fe- males in this small sample. Minor variations in color pattern are exhibited by the type material. The extremes in variation are exhibited by the primary types with the male being the darkest specimen collected and the female the lightest. Habitat.—The type series was collected from the margin of a pool in the bedrock of a small temporary stream. The stream on either side of the pool was dry at the time of collection as were many of the neighboring streams. Grass and brush grew at the stream margin and extended partially into the VOLUME 92, NUMBER 2 291 water. Most of the specimens were washed out of the roots and sand near these plants. Type data.—The Holotype, allotype and 11 paratypes were collected from a small unnamed stream where it crosses federal highway 59 approximately 2 miles south of Mena, Arkansas on July 19, 1978, by J. F. Matta. Deposition of type material.—The primary types are retained in the au- thors collection (JFM) but will eventually be deposited in the USNM and have been assigned type number 97566. Two paratypes are deposited in the NMNH and two paratypes are deposited in the CNC. The remaining para- types are retained by the authors; 4 by JFM and 3 by GWW. Etymology.—This species is named for the Ouachita mountains in which it was first collected. Hydroporus alleghenianus, new species Diagnosis.—A delicate little species which is similar to Hydroporus lae- tus Leech and Hydroporus cocheconis Fall and will key to the latter species in Fall’s (1923) key. It is smaller than either species with a maximum length of 3.1 mm. In addition it may be separated from cocheconis by the more distinct elytral maculation (the distal pale patches of the elytra are reduced in cocheconis) and from laetus by the pale discal and lateral portions of the pronotum. The aedeagus is similar to /aetus but has both the subapical tooth and the tip more distinctly produced ventrally. Description.—Holotype, male: Length 2.8 mm; width 1.35 mm; L/W = 2.07. Body form an elongate oval; elytra parallel sided for the basal one- third. Clypeal margin not thickened. Pronotum margined laterally, the mar- gin half as wide as the terminal antennal segment (approximately 0.1 mm), tapering slightly basally. Prosternal setae absent; prosternal process with a distinct protuberance and a reduced anterior file. The apical portion of the prosternal process almost flat, lanceolate shaped with a barely perceptible median raised area and with margins laterally. Head yellow brown with faint darker markings at the antennal bases. Antennae and palps a slightly darker yellow but without infuscation. Prono- tum yellow-brown, with a dark bar on the apical margin which does not reach beyond the eyes laterally. Elytra yellow-brown, lighter than the pronotum, with a brown stripe along the suture, a short brown stripe along the basal margin extending from the sutural stripe one-half the width of the elytra, a brown patch extending from the suture to the elytral margin sub- basally and another postmedially. The tips of the elytra are narrowly mar- gined with brown. The ventral surface light brown throughout, somewhat darkened at points of articulation. Head with surface microreticulate and finely punctured (about as in pul- cher). Pronotal punctation on the apical margin slightly coarser than the 292 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON head; becoming coarser basally and laterally; microreticulations present be- tween the punctures. Elytral punctation fine, with microreculations between the punctures. Elytra with a sparse covering of light setae. Ventral surface with the metacoxae and metasternum coarsely punctured. The abdomen finely punctured. Protarsi not enlarged, equipped with glandular pubescence ventrally; the claws almost identical, the anterior one slightly thickened. The aedeagus is distinctive with a strong subapical tooth and a rounded tip which is split dorsoventrally and also produced ventrally (Fig. Ic, d). Allotype.—Female, length 2.8 mm, width 1.4mm, L/W = 2.00. The female is similar to the male except as follows: slightly broader than the male with the ground color of the elytra slightly more flavous. The anterior pronotal spot is broader and more diffuse than in the male. Variation.—Average length 2.77 mm (2.6—3.1); average width 1.3 mm (1.2-1.5); L/W = 2.13; (n = 10). Specimens from Tennessee have the ground color of the head, pronotum and elytra darker—approaching reddish brown. Specimens from Connecticut have the dark markings of the prono- tum and elytra expanded; in some cases greatly so. The subbasal and post median dark spots are frequently joined and occasionally form a large dark spot covering most of the elytra. The basal dark area is rarely expanded and connected to the subbasal dark spot. The pronotal coloring is sometimes expanded and may cover the entire pronotum in rare instances. Habitat.—Rarely encountered throughout the east but taken in abundance where found. The primary types were taken at the margins of a small trib- utary of Back Creek near the point where it crosses Virginia Rt. 39. During the first collection (1973) the stream was flowing at a slow rate; during the second collection there was insufficient water to create a flow. During both collections H. alleghenianus were removed from the sandy margins of the stream. The beetles were observed swimming actively at the stream margin and hiding under small pieces of gravel. They rarely rested in the open, usually digging their way under small rocks or gravel before resting. Other aquatic beetles taken in the same collection include Hydroporus blanchardi Sherman, Hydroporus mellitus LeConte, Hydroporus stratiopunctatus Mel- shiemer, Hydroporus sulcipennis Fall, Hydroporus carolinus Fall, Hydro- porus pulcher LeConte, Uvarus sp., Paracymus confusus Wooldridge, Lac- cobius reflexipenis Malcolm and Dineutus discolor Aube. Type data.—The Holotype, allotype and 9 paratypes are from Bath Co., Virginia VIII-9-1973, J. F. Matta. Other paratypes (15) from the same lo- cality were collected VIII-7-77. Additional paratypes are designated as fol- lows: TENNESSEE, Morgan Co. Flat Fork Creek GWW+JAW, 2 August 1978, 53 specimens; Stewart Co. Hickman Creek, GWW+RER, 25 May 1978, 4 specimens. Etymology.—This species is named after the Alleghenian Section. VOLUME 92, NUMBER 2 | 293 Deposition of type material.—The primary types are retained in the au- thor’s collection (JFM) but will eventually be deposited in the (NMNH) and have been assigned type number 79567. Two paratypes have been placed in the NMNH, 2 in the Canadian National Collection and the remaining paratypes are retained by the authors. Literature Cited Fall, H. C. 1923. A revision of the North American species of Hydroporus and Agaporus.— Mt. Vernon, New York (J. D. Sherman, Jr.), 129 pp. Leech, H. B. 1949. Some Neartic species of hydradephagid water beetles new and old (co- leoptera).—Can. Ent. 80:89—96. Wolfe, G. W. and J. F. Matta. 1978. Three new species of Hydroporus (Coleoptera: Dytis- cidae) from the Southeastern United States.—Proc. Ent. Soc. Wash. (in press). (JFM) Dept. of Biological Sciences, Old Dominion University, Norfolk, Va. 23508; (GWW) Dept. of Zoology, University of Tennessee, Knoxville, Tennessee 37916. PROC. BIOL. SOC. WASH. 92(2), 1979, pp. 294-306 A NEW SPECIES OF CTENOCHELES (CRUSTACEA: DECAPODA: THALASSINIDEA) FROM THE NORTHWESTERN GULF OF MEXICO Nancy N. Rabalais! Abstract.—During an extensive U.S. Bureau of Land Management spon- sored survey of the south Texas outer continental shelf in the northwestern Gulf of Mexico, 5 specimens of a new species of Ctenocheles Kishinouye, 1926, were collected. The species, C. leviceps, is described and illustrated with a brief discussion of its ecology and occurrence. Introduction The genus Ctenocheles Kishinouye, 1926, first described from the Indo- West Pacific, consists of three species known from that region (C. balssi Kishinouye, 1926; C. collini Ward, 1945; and C. maorianus Powell, 1949) and representatives reported from the Atlantic Ocean. The first of these latter occurrences was based on major chelipeds of two species collected off Florida, Panama, and Columbia, which Holthuis (1967), for lack of com- plete material, designated as Ctenocheles A and B. The second was based on major chelipeds collected from Gabon, which Crosnier (1969) was un- willing to designate specifically. Specimens of C. serrifrons Le Loeuff and Intes, 1974, were collected from the continental shelf off Ivory Coast and described. Most recently, a new species from the western Atlantic, C. hol- thuisi Rodrigues, 1978, was reported off the mouth of Rio Sao Francisco, Brazil. During a U.S. Bureau of Land Management (BLM) sponsored survey of the south Texas outer continental shelf in the northwestern Gulf of Mexico, benthic ecologists from the University of Texas Port Aransas Marine Lab- oratory collected five specimens of a new Ctenocheles. Three specimens were collected in macroinfaunal (>0.5 mm) samples taken with a 0.025-m? Smith-McIntyre bottom grab; the others were taken with epifauna in a 10.7- m otter trawl. Samples were taken in 1976 and 1977 at 25 stations (1-6, 7) along 4 tran- sects (I-IV) situated perpendicular to shore from 10 to 130 m (Fig. 1). Ad- ditional stations included natural topographic high features at Hospital Rock (HR) and Southern Bank (SB), a rig monitoring station (RM), and a replicate trawling transect (TL) from which the holotype was collected. Detailed de- scription of the study area is given by Groover (1977). ' University of Texas Marine Science Institute Contribution No. 314. VOLUME 92, NUMBER 2 295 Ctenocheles leviceps, new species Figs. 2—29 Holotype.—Immature 3; complete specimen; total length (approximate), 34 mm; carapace length, 9.5 mm; Gulf of Mexico, ESE of Port Aransas, Texas; 27°38'N, 96°41’W (approximate); type-locality (TL) in Fig. 1; 40 m; 5 December 1977; R/V Longhorn; BLM night epifaunal sample; USNM 171576. Paratypes.—1 immature 6; complete specimen; total length (approxi- mate), 27 mm; carapace length, 6.1 mm; Gulf of Mexico, ESE of Port Aran- sas, Texas; 27°45'N, 96°43’W; 31 m; 2 March 1977; R/V Longhorn; BLM infaunal sample at RM; USNM 171577. 1 juvenile; complete body but no chelipeds; total length (approximate), 15 mm; carapace length, 3.8 mm; Gulf of Mexico, SE of Port Aransas, Texas; 27°30'N, 96°45'W; 49 m; 6 August 1976; R/V Longhorn; BLM in- faunal sample, Station 2, Transect II, Replicate 3; USNM 171578. 1 mutilated carapace and major cheliped; carapace length, 4.4mm; Gulf of Mexico, SE of Port Aransas, Texas; 27°30'N, 96°45’W; 47 m; 7 October 1976; R/V Longhorn; BLM infaunal sample, Station 2, Transect II, Repli- cate 1; USNM 171579. 1 mutilated carapace and minor cheliped; carapace length, 7.0 mm; Gulf of Mexico, S of Port O’Connor; 28°14'N, 96°29’'W (approximate); 10 m; 12 June 1976; R/V Longhorn; BLM night epifaunal sample, Station 4, Tran- sect I; USNM 171580. Diagnosis.—Major chela with elongate fingers armed on cutting edge with long, curved, acuminate teeth alternating with numbers of similar but small- er teeth. Palm shorter than fingers, 0.3 of total manus length, swollen, but not exaggerated. Minor cheliped, long and slender, relative lengths of arti- cles similar to major cheliped but without elongate, sharply toothed fingers. Fifth pereopods subchelate. Third maxilliped narrow, pediform with ele- vated, sharply toothed, thick crista dentata on distal *%4 to %4 of ventral surface. Telson slightly shorter than uropods. Posterior margin of uropodal exopod armed with acute spines. Description.—Carapace contained 3.55 times in total length of body, not covering whole length of cephalothorax, but leaving the last pereopodal segment entirely uncovered. Lateral margin ciliated. Carapace without well- defined oval. Linea thalassinica and cervical groove pronounced. Rostrum acute, not spinous, laterally compressed, with median dorsal carina and median ventral keel, dorsal margin straight, tip slightly deflected. Rostrum not reaching to end of eyestalks. Two tufts of small setae imme- diately behind rostral base. Median rostral carina continued posteriorly less than 4 length of carapace. Postrostral carina slightly elevated on gastric region, more pronounced elevation on cardiac region. 296 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 95°30’ —_— NAUTICAL ore a KILOMETERS. ea 0 20 40 60 ew fe CORPUS® ae) CHRISTI “;; OF MEXICO ¢ STUDY AREA \ PORT ISABEL Fig. 1. Map of nearshore northwestern Gulf of Mexico showing sampling stations and tran- sects and localities where Ctenocheles leviceps was collected. Eyestalks generally flattened anteriorly except for bulbous, faintly pig- mented subterminal cornea; longer (1.5 times) than wide, not reaching distal margin of first segment of antennular peduncle. Antennal peduncle reaching beyond antennular peduncle. Third antennular segment 1.7 times length of second. Anterior margins of third antennular segment and third antennal 297 VOLUME 92, NUMBER 2 “WU = J[BOG “MOIA [R1O}e] “ P sinjyewut ‘adAjojoy ‘sdadiaa] sajayoouayaD ‘7 BI 298 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Figs. 3-4. Ctenocheles leviceps, carapace: 3, Lateral view; 4, Dorsal view. Scale = 1 mm. segment at approximately same level. Third segment of antennal peduncle 1.4 times length of fourth. Two proximal segments of antennal peduncle equal in length to first antennular segment. Antennal scale spined. Following descriptions based on mouthparts dissected from one of para- types (USNM 171578). Mandible heavily chitinized, cup-shaped, proximal cutting edge armed with 3 teeth. Anterior edge armed with 2 larger teeth. Small tooth inside cupped area proximally. Mandibular palp 3-segmented, bearing short, stiff terminal hairs. Palp directed posteroventrally over man- dible. First maxilla (maxillula) with 2-segmented endopodite, distal edges of basis and coxa lined with short, spiniform setae. Endopodite of second maxilla, long and digitiform with slightly hooked tip. Scaphognathite enlarged, elongate, anterior lobe rounded distally, pos- terior lobe quadrate, flared. Endites lined with stiff, finely serrated setae. First maxilliped with oval endite bearing stiff finely-serrate setae mesially, narrow curved bladelike exopod, elongate epipod produced anteriorly in a short, narrow, quadrate lobe and posteriorly into an enlarged subquadrate projection, and short, truncate endopodite. Second maxilliped with short digitiform exopod. Dactylus, propodus, and —> Figs. 5-14. Ctenocheles leviceps, mouthparts, drawn from USNM 171578 unless otherwise indicated: 5, Mandible, lateral surface; 6, Mandible, inner surface; 7, First maxilla; 8, Second maxilla; 9, First maxilliped; 10, Second maxilliped; 11, Second maxilliped, enlarged dactyl; 12, VOLUME 92, NUMBER 2 299 Third maxilliped, mesial surface, USNM 171579; 13, Third maxilliped, outline without setae, USNM 171579; 14, Third maxilliped, crista dentata, holotype. 5, 6, 8-10, scale = .5 mm; 7, 11, scale = .25 mm; 12-14, scale = 1 mm. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 300 VOLUME 92, NUMBER 2 301 carpus of equal width; propodus longer than dactylus and carpus of equal length. Merus slightly curved and longer than first 3 articles combined. Dactylus bearing distal setae armed with double row of blunt teeth. Third maxilliped narrow, pediform, with exopod. Merus and ischium com- bined length 3.5 to 4.5 greatest width. Elevated, sharply toothed, thick crista dentata on distal 24 to %4 of ventral surface of ischium. Teeth broadly tri- angular, not spinous. Carpus enlarged distally, with dense tuft of stiff setae on anteromesial surface. Propodus elongate, length 2.0 times width, with dense tuft of stiff setae on mid-lower mesial surface. Dactylus narrowest and shortest of articles, rounded distally. First pereopods unequal in size and form. Right cheliped longer (1.7 times) and more developed than left. Major cheliped 0.7 times total body length. Fingers of major chela slender, 1.8 times palm length, ending in blunt, crossed, hook-like tips, dactylus to inside. Fixed finger laterally compressed throughout length, height uniformly decreasing distally. Dactylus broadly rounded at base where attached to palm, becoming laterally compressed distally; height greater than fixed finger, particularly in distal half. Cutting edge of both fingers armed with long, curved, acuminate teeth alternating with numbers of similar but smaller teeth. Longest tooth on dactylus, 0.5 mm. Arrangement of long with shorter teeth similar to that described for Ctenocheles A and B-by Holthuis (1967). Palm shorter than fingers, 0.3 times length of whole hand, and swollen but not exaggerated; width greater than combined width of closed fingers; upper anterior surface flattened, remainder laterally compressed, equally convex on both sides. Carpus very short, articulating with palm along its anterolateral border as if fused; rough- ly triangular with rounded ventral border but dorsal margin straighter, more compressed, forming a bridge; articulating with merus on mesial surface at its posterodorsal edge. Merus elongate, tapering at both ends, length 4.0 times width. Ischium longer than merus, narrow (length 7.0 times width), and slightly sigmoid. Other than teeth on fingers, major cheliped is un- toothed and unspined, with sparse, short setae on dactylus, remainder smooth and mostly naked. Major cheliped of smaller specimens more se- tose. Minor cheliped long, slender, relative lengths of articles similar to major cheliped, but without elongate, sharply toothed fingers. Fingers slightly lon- << Figs. 15-24. Ctenocheles leviceps, pereopods, drawn from holotype unless otherwise in- dicated: 15, Right pereopod 1, lateral surface; 16, Right pereopod 1, lateral surface, USNM 171579; 17, Right pereopod 1, mesial surface, USNM 171579; 18, Left pereopod 1, lateral sur- face; 19, Left pereopod 1, lateral surface, USNM 171577; 20, Right pereopod 2; 21, Left pere- opod 3; 22, Left pereopod 4; 23, Right pereopod 5, lateral surface; 24, Right pereopod 5, en- larged mesial view of dactylus and propodus, 15—23, scale = 2 mm; 24, scale = 1 mm. 302 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON ger (1.2 times) than palm, laterally compressed, and tapered uniformly to bluntly pointed, slightly hooked, crossed tips; cutting edges slightly serrate. Smaller specimens without serrations on minor chela. Palm slightly wider than combined width of closed fingers, straight on dorsal and ventral mar- gins; articulating with carpus on entire anterolateral margin, similar to major cheliped. Carpus short, elongate-triangular, tapering proximally; ventral surface rounded, dorsal straight. Merus and ischium equal in length, both 4.0 times longer than wide; ischium narrowest in proximal third. Minor cheliped untoothed and unspined, with sparse, short setae on fingers, palm and carpus, remainder smooth and naked. Second pereopods chelate, compressed. Fingers 2.0 times palm length, cutting edges smooth and sharp. Manus with numerous tufts of short setae. Carpus slightly shorter than manus. Merus length 1.7 times carpus and 4.8 times ischium. Long setae present on ventral margins of ischium and merus, anterodorsal edge of merus, dorsal margin of carpus, and anteroventral edge and anterior border of carpus. Third pereopods as long as second. Dactylus twice as long as wide, acute- ly pointed. Propodus and distal part of carpus wide (4.0 times widest part of dactylus). Dactylus and propodus bearing long, fine setae on dorsal and ventral margins, dorsolateral areas of the propodus setose, and anteroven- tral region of carpus densely setose with long, fine hairs and stiff, short setae. Merus 1.4 times as long as carpus and 3.2 times as long as wide. Fourth pereopods slightly longer and more slender than third. Dactylus long, slender, acute; propodus wide; carpus long, club-shaped. Dorsal and ventral margins of dactylus, propodus and anterior end of carpus bearing long, fine hairs; propodus setose on dorsolateral edge with dense tuft of stiff, short setae distally on ventrolateral edge. Relative lengths of articles similar to third pereopods. Fifth pereopods long, narrow and subchelate. Dactylus elongate, curved and flattened, twisting inward and downward halfway toward the distal end, ending in flattened, triangular flare. Fixed finger blunt, short and spoon- shaped. Dactylus length 2.0 times fixed finger. Propodus uniformly curved, 4.4 times longer than wide. Transverse ridge crossing propodus from mid- ventral to anterodorsal borders supporting dense row of thick, stiff setae. Otherwise, dorsal and ventral margins of propodus and dactylus lined with fine, long setae. Carpus and merus scantily haired. Carpus uniformly curved, widened distally. Arthrobranchs present from third maxilliped to fourth pereopods, inclu- sive, numbering 2 per appendage. Abdomen much elongated, pleura little developed and smooth. First ab- dominal segment widest (1.2 times _ ider than long); sixth segment longest (1.2 times longer than second, 1.5 times longer than third, 1.6 times longer than fourth, and 2 times longer than fifth). Dorsal surface of each pleuron VOLUME 92, NUMBER 2 303 S 4 Y 60 = 7) tp 40 © zoea 2 Pe) Cc ® 20 1S) hon Pd megalopa 2 4 6 8 10 12 14 16 18 Survival in days VOLUME 92, NUMBER 2 311 curred and had already molted to zoeal stage I by the time the larvae were examined. The prezoeal molt is especially diaphanous and could easily have been overlooked amid the debris of egg casts on the bottom of the covered glass hatching bowl. The duration of each development stage is presented in Table 1, and the percentage of larval survival is illustrated in Fig. 1. Although some variation in developmental time was seen among the fed series at different tempera- tures, in general it required at least 14 days (at 25°C), and 16 days (at 20°C) to attain crab stage I. Larvae usually remained in stage I from 3-4 days at all temperatures although some zoeae held at 20°C often required 5 days to reach stage II. Second stage zoeae usually remained as such for 3-4 days at 25°C and 3-5 days at 20°C, so little temporal variation occurred within this instar. Based on rather limited data of two surviving instars the duration of the megalopal stage was also consistent, lasting 8—9 days at both of these temperatures. The duration of development of larvae in the diel-pro- grammed CTU at 25°C paralleled that seen at warm room temperature, but with a slightly shorter second zoeal stage (2 days). No conclusions can be drawn regarding the postlarvae duration because both surviving megalopae died after one and two days in stage, respectively. Larvae in the 30°C CTU did not survive through the second zoeal stage, and all starved zoeae died in stage I. ¥ These results suggest that M. forceps can complete its planktonic exis- tence in a period approximating two weeks. This is at variance with the developmental time noted by Provenzano and Brownell (1977) for M. spi- nosissimus which required the relatively short duration of 130-148 hours (5-6 days) to attain first crab stage at temperatures of 24—28°C. Whether this rapid development is more a result of the variable, higher temperatures in their program, or is inherent in the larvae of their species, or perhaps both, cannot be ascertained at present. For our part, we did note a more rapid development in stage II zoeae held in the diel fluctuating temperatures of 25°-20°C so that the megalopal stage was reached after a total zoeal duration of about 5 days (120+ hours). Comparatively, megalopa was reached in M. spinosissimus in only 2—3 days (60-70 hours), but because none of our megalopae in the diel program survived, further speculations are unwarranted. However, the results of our differential culturing-temper- ature program show, as seen in other laboratory-culture studies on decapod larvae, that temperature affects duration of development, with cooler tem- peratures (i.e. 20°C) increasing developmental time, and warmer tempera- tures (i.e. 25°C) decreasing it (see Fig. 1). << Fig. 1. Mithrax forceps: Percentage and duration of survival of larvae cultured under lab- oratory conditions. See text for explanation of diel variation. 312 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Description of Developmental Stages First Zoea.—(Carapace length.—70 mm; number of specimens examined, 10). Carapace (Fig. 2A, a): Cephalothorax smooth, inflated, globose; with short recurved dorsal and straight rostral spine. A dorsal tubercle medially, midway between the bases of dorsal and rostral spines, followed by a pair of minute setae postero-laterally, and a second pair dorso-laterally on car- diac region. Six setae on posterolateral border, first (=anterior seta, Yang 1967) by far the strongest (Fig. 2a). Thoracic appendages unsegmented, extending slightly below posterolateral margin of carapace. Eyes unstalked. Abdomen (Fig. 2A): Five somites, all with pair of small setae postero- dorsally; second with pair of small lateral spines curving anteriorly; third through fifth each with a single short spine on posteroventral angle. Pleopod primordia on second through fifth somites amorphous, subdivided. Telson (Fig. 2B): Trapezoidal, not deeply excavated posteriorly, but with elongate furcae; latter covered with small fine hairs and carrying a pair of lateral movable spines at base; posterior margin of telson with 6 stout setae, armed with rows of spinules. Antennule (Fig. 2C): A conical rod; 2 long stout, and 2 thinner, short terminal aesthetascs, plus | fine hair. Antenna (Fig. 2D): Protopodite a slender dagger-like process bearing 2 rows of small teeth from midlength distally to tip. Exopodite tapered, equal to or slightly longer than protopodite; distal quarter with 2 rows of small teeth; a slender serrated spine and a naked seta distally. Endopodite bud naked, 44 length of protopodite. Mandible (Fig. 2E): Asymmetrically dentate, scoop-shaped processes. Left incisor margin with 5 distinctly rounded prominences; molar irregularly dentate; 3 rounded prominences at junction of molar and incisor of left side (ventral view); right side with 1 prominence. Maxillule (Fig. 2F): Endopodite 2-segmented; proximal short, with 1 long feathery seta laterally, distal longer, with 4 unequal terminal setae plus 2 longer setae subterminally. Coxal and basal endite each with 7 terminal processes, 5 of which on basal endite are stouter. Maxilla (Fig. 2G): Endopodite unsegmented, 5 terminal setae. Proximal and distal lobes of coxal and basal endites each with 5, 4 setae, respectively; pubescence on basal endite and endopodite as illustrated. Scaphognathite with 13 plumose setae on outer margin, distal setae increasing in stoutness, as in detail (Fig. 2g). — Fig. 2. Mithrax forceps, first zoea: A, Lateral view; a, Detail of posterolateral margin of carapace; B, Telson; C, Antennule; D, Antenna; E, Mandibles; F, Maxillule; G, Maxilla; g, Detail of scaphognathite apex; H, Maxilliped 1; I, Maxilliped 2. VOLUME 92, NUMBER 2 313 314 A-B PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 1.0mm C-J 0.25mm /——— VOLUME 92, NUMBER 2 3145 Maxilliped 1 (Fig. 2H): Coxopodite with | seta. Basipodite with 10 setae, progressing distally, 2, 2, 3, 3. Endopodite 5-segmented, ventral setal for- mula progressing distally 3, 2, 1, 2, 4 + I (roman numeral denotes dorsal seta). Exopodite incompletely 2-segmented, 4 terminal natatory setae. Maxilliped 2 (Fig. 21): Coxopodite naked. Basipodite with 3 ventral setae. Endopodite 3-segmented, ventral setal formula progressing distally 0, 1, 5; heaviest seta placed laterally. Exopodite as in maxilliped 1. Color: Zoea transparent overall; frontal region, gut region and dorsal sur- face of abdominal somites pale gold; abdominal somites 2, 3, 4, orange ventrally, fifth only faintly so. Several spider-like black chromatophores on labrum and mandible; a line of same extends interiorly from gut through second abdominal somite, spreading only slightly into third. Gut green in- teriorly. Eyes turquoise in reflected light, corneas dark. Second Zoea.—(Carapace length.—0.82 mm; number of specimens exam- ined, 10). Carapace (Fig. 3A): Cephalothorax essentially unchanged in general form from stage I, but larger, more inflated, dorsal spine proportionately shorter. Paired dorsal setae now include 2 interocular, 2 posterolateral to dorsal tubercle, 1 at each lateral base of dorsal spine, plus pair previously noted dorsolaterally on cardiac region. Carapacial setation on posterolateral mar- gin now | strong, 6 finer. Thoracic appendages longer but remain unseg- mented. Eyes stalked, a minute ocular tubercle on anterior margin. Abdomen (Fig. 3A): Six somites; first with 3 setae on posterodorsal mar- gin, second with one median and one posterodorsal pair, third through fifth unchanged from stage I. Lateral spine on somite 2 unchanged, spines pos- teroventrally on somites 3—5 more elongate than in stage I; sixth somite now with single spine on posteroventral angle. Pleopod buds well-developed on somites 2—5, rudimentary on somite 6. Telson (Fig. 3B): Similar in form and marginal setation to first stage. Furcae same length as stage I but now 3 times length of reduced telson, maintaining posterior spines and hairs. Antennule (Fig. 3C): Endopodite bud present, placed as shown. Seven unequal aesthetascs and single fine hair terminally. Antenna (Fig. 3D): Similar in form and armature to first stage. Endopodal bud lengthened, now nearly % protopodite process. Mandible (Fig. 3E): As in first stage; palp bud present on each anterior surface. Maxillule (Fig. 3F): Endopodite unchanged. Coxal endite with 7, basal <_< Fig. 3. Mithrax forceps, second zoea: A, Lateral view; B, Telson; C, Antennule; D, An- tenna; E, Mandibles; F, Maxillule; G, Maxilla; H, Maxilliped 1; I, Maxilliped 2; J, Maxilli- ped 3. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 316 VOLUME 92, NUMBER 2 ay endite with 10, processes of varying stoutness; a single seta dorsally on lower outer margin of basal endite. Maxilla (Fig. 3G): Endopodite and coxal endites unchanged in form and setation from first stage, basal endite now with 5 setae on each lobe. Sca- phognathite with 24 plumose marginal setae. Maxilliped 1 and 2 (Figs. 3H, I): Similar to first stage; exopodite on each with 6 terminal natatory setae. Maxilliped 3 (Fig. 3J): Naked, bilobed, rudimentary process. Color: Cephalothorax and abdominal coloration similar to first stage. Po- sition and color of chromatophores essentially unchanged except for addi- tional black chromatophore on carapace posterio-dorsad to each eye; chro- matophores on gut and maxilliped 2 more pronounced, that on maxilliped 1 now lacking. Eye color similar to stage I. Megalopa.—(Carapace length x width.—1.06 x 0.96 mm; number of specimens examined; 10). Carapace (Fig. 4A, B): Cephalothorax generally ovally subquadrate, with well developed supraocular lobes, outer orbital angles bluntly produced; frontal region rectangular, rostrum elongate, deflexed, spiniform. Gastric region with 3 large transverse tubercles, plus a pair immediately behind outer 2; cardiac region with 5 small tubercles forming an arch. Single, largest tubercle on intestinal region. Eyes large, anterior margin of eyestalks with 2 small setae. Abdomen (Fig. 4A—B, 5A—C): Abdominal pleura 2—5 with lobes at pos- teroventral angles, that of sixth subquadrate, all with minor setation placed as shown. Telson smooth, subquadrate, posterior angles rounded, 2 dor- somedial setae. Pleopods of decreasing size on somites 2-6, with 11, 11, 11, 9, 5 setae on each exopodite, respectively. Endopodites on pleopods 1-4 without setae but with appendix interna; pleopod 5 (=uropod) without en- dopod or appendix interna. Antennule (Fig. 4C): Biramous. Peduncle partly or completely 3-seg- mented; basal segment naked, second with 2, third with 1 distal, setae. Lower ramus with 3 setae, placed 2 terminally, 1 subterminally; upper ramus 2- or incompletely 3-segmented; penultimate with 7 aesthetascs placed in V, progressing distally 2, 2, 2, 1, plus 1 short distoventral seta; terminal seg- ment with 5 aesthetascs, progressing distally as 4, 1. Antenna (Fig. 4D): Peduncle with 2 distolateral lobes and a single seta. Setation of the 6 flagellar segments proceeding distally 2, 3, 0, 0, 4, 3, plus 1 hair. oe Fig. 4. Mithrax forceps, megalopa, sensory and feeding appendages: A, Lateral view; B, Dorsal view; C, Antennule; D, Antenna; E, Mandibles; F, Maxillule; G, Maxilla; H, Maxilliped 1; I, Maxilliped 2; J, Maxilliped 3. 318 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Mandible (Fig. 4E): Incisor now truncately spatulate, margin heavily chi- tinized. Palp indistinctly 2-segmented, with 0, 5 setae, respectively. Molar process irregularly serrate. Maxillule (Fig. 4F): Endopodite indistinctly 2-segmented, naked. Coxal endite with 10, basal endite with 18, marginal processes of varying stout- ness; latter retains 1 long feathery seta on lower margin. Maxilla (Fig. 4G): Endopodite inflated, unsegmented, naked. Setation on proximal and distal lobes of coxal and basal endites 7, 3, and 6, 6, respec- tively. Scaphognathite with 26-30 setae on outer margin, plus 3 laterally on either side of blade. Maxilliped | (Fig. 4H): Endopodite partly 2-segmented, naked. Exopodite 2-segmented, setation 1, 4, respectively. Coxal endite with 5-7, basal endite 9-11, marginal setae. Epipodite with 1 proximal, and up to 5 distal, setae. Maxilliped 2 (Fig. 41): Endopodite 4-segmented; setation progressing dis- tally 0, 1, 3, 6. Exopodite 2-segmented, 4 terminal setae. Epipodite absent. Maxilliped 3 (Fig. 4J): Endopodite 5-segmented, setal formula progressing distally usually 12, 9, 5, 4-6, 3-4. Exopodite 2-segmented, 6 terminal setae. Epipodite usually with 5, rarely 6, setae in illustrated positions. Protopodite ringed with 5—7 short setae. Pereiopods (Fig. SD—F): Chelipeds large, elongate, equal, setose. Manus smooth, inner and outer surface with scattered hairs; propodus with several small teeth distally on cutting edge, tip slightly hooked, overlapping dactyl exteriorly when closed; dactyl smooth, curved, setae as shown. Pereiopods 2—5 (walking legs 1—4) similar in form, setation sparse to moderate, as il- lustrated, all without supernumerary teeth on dactyls; as in other mayid megalopae pereiopod 5 lacking ““brachyuran feelers.’’ Color: Carapace overall a pale golden-brown; frontal region pale greenish- gold. Gut interiorly iridescent green, with paired black chromatophores dor- sally and posterolaterally. Mandible, labrum and protopodite of third max- illiped with several interlocking black chromatophores dorsally and pos- terolaterally. Coxopodite of cheliped with one black, one red-orange chromatophore ventrally; coxopodite and carpus of second through fifth pereiopods with a single pale red chromatophore ventrally. A single black chromatophore dorsally on anterior margin of abdominal somite 1; several spider-like red chromatophores at junction of somites 2-3; numerous, interspersed red and black chromatophores similarly placed at junctions of somites 3—4, 4-5, 5—6. Junction of somite 6 and telson pale greenish-gold, overlain with several small spidery black chromato- = Fig. 5. Mithrax forceps, megalopal locomotory appendages: A—C, Pleopods 1, 4, 5; D-F, Pereiopods 1, 3, 5. 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Telson anteriorly with a single golden-orange chromatophore me- dially. Peripheral ommatidia of eyes reflect blue light, corneas dark with pink highlights. Discussion Only limited comparison can be made among the presently known larvae of three species of Mithrax. Our data indicate that the zoeae of M. (Mith- raculus) forceps are much closer morphologically to at least the first zoeal stage of M. (Mithrax) pleuracanthus (based on features described by Yang, 1967), than they are to larvae of M. (Mithrax) spinosissimus (Provenzano and Brownell, 1977). The similarities are summarized in Table 2 where it can be seen that the only salient differences in morphological features be- tween the first zoeal stages of M. forceps and M. pleuracanthus are in number of antennular aesthetascs (4 and 3, respectively [but only 2 illus- trated by Yang]), the setal formula of the endopodite of maxilliped 2 (0, 1, 5 and 0, 1, 4, respectively), and in the admittedly subjective observation on the presence (M. forceps) or absence (M. pleuracanthus) of pleopodal pri- mordia. Other carapacial features such as the reach of the rostral spine of the antennule, and the robustness and curvature of the dorsal spine, are not now useful in distinguishing Mithrax larvae because of their subjectivity. The larvae of M. forceps and the first zoeal stage of M. pleuracanthus both differ greatly from the zoeal stages of M. spinosissimus. The zoeae of the latter species are very unusual in that several appendages, such as the maxillulary and maxillary endites, and the endopodites of both maxillipeds, exhibit a reduction in setation as compared to no such reduction in the former species. Such reduction in setal number, in conjunction with a lack of segmentation in some appendages (most notably in the endopodites of maxilliped 2) may be features indicative of an advanced stage in the larvae of M. spinosissimus, if decreasing morphological complexity is suggestive of apomorphy as espoused by Lebour (1928). Whether such apomorphy (if real) is characteristic of any other species in the subgenus Mithrax (sensu lato) remains to be determined. These reduced numbers are evident from the data in Table 2. The general form of the telson in the larvae of M. forceps and M. pleu- racanthus is relatively similar to that seen among the zoeae of other Mith- racine species. However, the larval telson of M. spinosissimus differs from M. pleuracanthus (in the same subgenus), and exhibits a form not close to the genus Mithrax, but rather to that seen in the Inachine genus Steno- rhynchus (see Yang, 1976). The differences in telsonal morphology are es- pecially notable in the median notch on the posterior margin, which, in M. spinosissimus (and in Stenorhynchus larvae) is very well developed, but is very much reduced in M. pleuracanthus, and almost totally absent in M. VOLUME 92, NUMBER 2 325 forceps. Indeed, the only telsonal feature shared with any degree of simi- larity between M. spinosissimus and the other two Mithrax is the setal formula on the posterior margin, which remains 3 + 3 in all larval stages. In comparing the zoeal features of M. forceps and M. spinosissimus we noted in Provenzano and Brownell’s illustrations (1977; Figs. 2B, 3A, B) the presence of 5 pairs of pleopod buds in stage I, and the apparent lack of a sixth abdominal somite in stage II. The last feature was especially intrigu- ing because, as far as is known, only larvae of Micippa in the Mithracinae exhibit 5 abdominal somites in both zoeal stages. With the exception of Menaethius (Acanthonychinae), and Achaeus, Inachus and Macropodia (Inachinae) all other presently known majid larvae have 6 somites in stage II. However, Provenzano (personal communication) clarified this apparent anomaly, stating that the fifth pair of pleopods in stage I (his Fig. 2B) are probably non-erupted pleopodal buds which may have been visible through the zoeal abdomen. A re-examination of stage I] using more powerful optics by Provenzano showed that a sixth somite was definitely present but the dividing suture between the telson and that somite was very fine, and the associated pleopod buds were extremely small and easily overlooked. Thus, M. spinosissimus exhibits 5 abdominal somites in stage I, and 6 in stage II, as seen in other Mithracinae larvae. Provenzano also informed us that a mandibular palp was present in stage II, thereby providing yet another fea- ture of agreement between the zoeae of M. spinosissimus and M. forceps, and consequently with other known Mithracine larvae. In regard to other larval features within the subfamily Mithracinae, or even more broadly considered, among the family Mayjidae, little can be said. We tend to agree with Provenzano and Brownell (1977) who stated that there are few characters which can be used to distinguish among the larvae of the various genera. We did note, as did Yang (1967) and Kurata (1969) that many of the genera exhibit a remarkable consistency in larval features, with exactly similar formulae in antennular aesthetascs, processes on max- illulary and maxillary endites, in basipodal, endopodal and exopodal seta- tion, and spination on abdominal somites. The chief differences among the larvae of the Western Atlantic genera appear to be in the setal formula on the posterolateral carapace margin, the scaphognathite, and in the general form of the telson. The difficulty in separating majid larvae is therefore probably more often exacerbated by consistency in zoeal features than not. It is obvious that many more studies within the genera are needed before the relative importance of most larval features can be finally determined. Data on Mithracine postlarval stages are even more sparse, sO we must confine our remarks to a comparison between M. forceps and M. spinosis- simus (Table 2). As might, perhaps, be expected in a taxon wherein the adult exhibit so much heterogeneity in form, the megalopae of the two 326 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON species can be easily distinguished by their respective overall morphology, although neither resembles in form a minature adult as is true for some anomuran and brachyuran postlarvae. The postlarval stages of M. spino- sissimus, like the zoeal stages, continued to show reduced or absent setation or segmentation on the mouthparts, in comparison to that seen in M. for- ceps. However, the latter species, like M. spinosissimus, now also lacks setae on some appendages, notably the endopodites of the maxillule, max- illa, and maxilliped 1. But the two species may otherwise be easily differ- entiated using antennular aesthetascs (more in M. forceps), scaphognathite marginal setae (less in M. forceps), second maxilliped endopodal segmen- tation (4 in M. forceps, 5 in M. spinosissimus), and general armature on maxilliped 3 (exopodite setose, endopodite ischium unarmed, a ring of setae on the basipodite in M. forceps; exopodite naked, endopodite ischium den- tate, no basipodal setal ring in M. spinosissimus). The reduced setation and segmentation in the megalopal appendages of M. spinosissimus reflect a situation noted earlier in the zoeal stages, again suggesting that if apomorphy has occurred the postlarval stage of this species may be more advanced than that of M. forceps. Larval and postlarval studies on M. pleuracanthus and related species should therefore prove most interesting in this respect. The very close similarity between the zoeae of M. forceps and M. pleu- racanthus raises some question as to the utility of their respective subge- neric groupings in view of the different larval features now known to occur. It is additionally apparent that the larvae of M. spinosissimus can be par- ticularly distinguished on morphological grounds from at least one species (i.e. M. pleuracanthus) presently occupying the same subgenus. Moreover, the data presented above show that M. spinosissimus is clearly less related in several zoeal and megalopal features to other members of the subfamily Mithracinae. Rathbun (1925) characterized the two subgenera primarily by the presence (Mithraculus) or absence (Mithrax) of branchial sulci in the adults, in con- junction with more subjective features such as the shape and length of ros- tral horns, and the development of minor teeth of the orbit. However, the very close morphological similarity seen in at least the first zoeal stages of two species which are separated subgenerically as adults primarily by the presence (M. forceps) or absence (M. pleuracanthus) of branchial sulci, and the inclusion in the subgenus Mithrax of the third species (M. spinosissimus) which differs not only from the former two species, but in many features from other Mithracine larvae as well, argues for a reassessment of the genus Mithrax, and certainly a redefinition of the particular subgeneric diagnoses to take into account recently delineated larval characters. For example, the genus could eventually be separated into two genera (perhaps by elevation of a subgenus after appropriate redefinition), or a third subgenus erected to VOLUME 92, NUMBER 2 327 accommodate those adults which exhibit larval features obviously disparate from other species. Acknowledgments We thank Messrs. John K. Reed and Frank G. Stanton, Harbor Branch Foundation, Inc., Link Port, Ft. Pierce, Florida, for collecting the ovigerous female specimens used in this study. We are extremely grateful to Dr. An- thony J. Provenzano, Jr., Institute of Oceanography, Old Dominion Uni- versity, Norfolk, Virginia, for graciously re-examining some of the speci- mens he used in his study, and for providing additional data on larval morphology at our request. Literature Cited Gore, R. H. 1971. Pachycheles monilifer (Dana, 1852): The development in the laboratory of larvae from an Atlantic specimen with a discussion of some larval characters in the genus (Crustacea: Decapoda; Anomura).—Biol. Bull., Woods Hole 144:132—150, figs. 1-6. Kurata, H. 1969. Larvae of Decapoda Brachyura of Arasaki, Sagami Bay—IV. Majidae.— Bull. Tokai Reg. Fish. Res. Lab., No. 57:81-127, figs. 1-27. Lebour, M. V. 1928. The larval stages of the Plymouth Brachyura.—Proc. Zool. Soc. Lond. 1928 (2):473-560, pls._1—16, text-figs. 1-5. . 1944. Larval crabs from Bermuda.—Zoologica, N.Y. 29:113—128, figs. 1-19. Provenzano, A. J., Jr., and W. N. Brownell. 1977. Larval and early post-larval stages of the West Indian spider crab, Mithrax spinosissimus (Lamarck) (Decapoda: Mayidae).— Proc. Biol. Soc. Wash. 90:735-752, figs. 1-7. Rathbun, M. J. 1925. The spider crabs of America.—Bull. U.S. Nat. Mus. 129:i-xx, 1-613, pls. 1-283, text-figs. 1-153. Williams, A. B. 1965. Marine Decapod Crustaceans of the Carolinas.—Fishery Bull. U.S. Fish Wildl. Serv. 65:i-ix, 1-298, text-figs. 1-252. Yang, W. T. 1967. A study of zoeal, megalopal and early crab stages of some oxyrhynchous crabs (Crustacea: Decapoda).—Ph.D. dissertation, University of Miami, Coral Gables, Florida, pp. i—xiii, 1-459, figs. 1-94. . 1976. Studies on the western Atlantic arrow crab genus Stenorhynchus (Decapoda Brachyura, Majidae) I. Larval characters of two species and comparison with other larvae of Inachinae.—Crustaceana 31:157—177, figs. 1-13. Smithsonian Institution, Fort Pierce Bureau, Ft. Pierce, Florida 33450. PROC. BIOL. SOC. WASH. 92(2), 1979, pp. 328-332 MESOSIGNUM ANTARCTICUM, NEW SPECIES, THE FIRST RECORD OF THE GENUS FROM THE DEEP SEA SOUTH OF THE ANTARCTIC CONVERGENCE (ISOPODA: JANIRIOIDEA) George A. Schultz Abstract.—A single female specimen of a new species of Mesosignum Menzies, 1962—M. antarcticum—was obtained from the deep sea south of the Antarctic Convergence in the Pacific Quadrant of the Antarctic Ocean. The species is described and illustrated and its place in the genus is dis- cussed. A single female specimen of Mesosignum Menzies (1962) was collected at a R/V Eltanin station in the deep part of the Pacific Quadrant of the Antarctic Ocean. It was the only specimen of the genus discovered from among thousands of specimens from more than 820 stations examined from the whole Antarctic region. It is the first of the genus to be taken far from the Pacific and Caribbean locations where other species of the genus have been recorded. Menzies (1962) based the genus on M. kohleri, a species which he described along with one other from the Caribbean Sea. Birstein (1963a) described two species from the north Pacific and later (1963b) one from the Bougainville Trench of the Pacific. Menzies and Frankenberg (1968) described six species from the central western Pacific and compared them to the previously described species. Menzies and George (1972) re- described one of the Menzies and Frankenberg species and described a new one from the Peru-Chile Trench. The total species in the genus including the new one described here is 13. Their length, sex or sexes known, depth and distribution are included in Table 1. Most species were collected in the deep sea within 30°N and 30°S of the equator. The depth range for the species of Mesosignum is from 1016-7954 m with most species being from below 3000 m. The species are blind and without pigment as are most other Janirioidea from the deep sea. Mesosignum antarcticum, new species Figs. 1-12 Description.—Cephalon and peraeonal segment I folded so most of dor- sum of cephalon not visible in dorsal view. Dorsum of cephalon and pe- VOLUME 92, NUMBER 2 329 Figs. 1-8. Mesosignum antarcticum: 1, Dorsal view holotype female 2.5 mm long; 2, An- tenna 1; 2, Maxilliped; 4, Mandible (damaged); 5, Mandible; 6, Operculate pleopod; 7, Pleopod 3; 8, Hypopharynx. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 330 yously, UleJLIg MON ‘oyloeg eluosejeg 0} BOLISUTY YINOG }SVODd ISOM eVIQUIO[OD ‘eudsozIeZ JO JSOMYJIOU—BISg UROQGqUe) qousry, sty D-Mled ROTY VISOD JJO dyIOeg eVOIY VISOD JJO dyIoeg uvd00O II}DIEJUY ‘JUeIPEN() dYIOeg BNSVILSIN JJO 9yIOeg Yous], sIyO-n1eg BIg UBSqqLIeD ‘osesseg PICMPUIAA Bag UvIGqUeD ‘uUIe[g URIquUIO[OD ueder ‘eyescC Jo unos Spuv[s] I[LINy JO yseoyjnos BOY BISOD JJO oylOeg uones07] vS6L—-0769 S90r—-9I0I SC6b-EC8P O9CE-PSTE SILt 6t0P OS6E-LLLE pSE9O-CLEE 8Ltt ILOb—8982 OST ¥—000PF 089S—0L9S 69€I—-9101 yideq P Oe oe Vc 5 (SG 6) 8°C ‘2 Oy Ik 6) sc } Vc Ow Oe fo) 97 Of 4 6) 61 6) 91 62 Ge uMOUY yj3u9] SOxos WINUIIXxe] IO X9S ee eee SO oasw—x—m—x— rrr eee QE96I ‘UlaISIIg 120/114 796 “SOIZUSP Maysn ZL6[ ‘981IOIH pur soizusp wnjnouny 896] ‘sloquoyuel, pue soizuay wnionw 896] ‘dIoquoyuely pue soizusp, wnsadsp 6L6I1 °ZI[NYIS wnd1I4DJUD 8961 ‘sloquoyuely pue soizuayy wnjvsun uos[e}0a|d Woy suUONdafo1d 3u0J—g od], 8961 ‘Sioquoyuely pue soizusyp, suapiyjniu 896] ‘sIoquoyuely pue soizusp, suappuspw (sa1sads-9dA}) 796] ‘SoIZUay Majyoy VEQ6] ‘UlOISIIQ Wnjnjudsaja VEO “UlOJSIIG SIUIdsi1Aa4g 8961 ‘sloquoyULIy pue soizusp, wnjosiuppo uos[a}09[d Woy sUONdafoid a31e] ON—VW odA], iii ———— ‘so1seds wnusisosaw “| AQey, VOLUME 92, NUMBER 2 35 Figs. 9-12. Mesosignum antarcticum: 9-11, Peraeopods I, II, and VII respectively; 12, Detail dactylus peraeopod II. raeonal segment I without dorsal or lateral spines. Dorsum of peraeonal segments II and III with elongate anterolateral corners and short pointed posterolateral corners. Peraeonal segment IV with elongate anterolateral corners of about half length of extensions on peraeonal segments III and V; without posterolateral extensions. Peraeonal’ segment VI with very long posteriorly directed lateral extensions; extensions longest of all peraeonal extensions and envelop most of length of pleotelson. Peraeonal segment VII about as wide as peraeonal segment I and without lateral extensions. Pleo- telson longer than broad with serrated, convex lateral borders; posterolater- al extensions of pleotelson slightly longer than anterolateral extensions on segment IV. Posterior margin of pleotelson convex. 332 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Antenna | with 7 segments; apical one with single aesthetac and one long seta. (Mouth parts damaged.) Mandible with lacinia mobilis; 4 setae in setal row. Molar process thin and elongate with long apical setae. Mandibular palps broken; basal palp article present on one mandible. Maxilliped with all palp articles narrow; 2 coupling hooks; endite with plain (probably dam- aged) sensory edge. All peraeopods long and thin with I the least and VII the most elongate. All dactyli with elongate unguis as major claw. Oper- culate pleopod ovate with 11 setae on convex posterior margin. Uropods short, not entending beyond posterior margin (dorsal view.) Type-locality.—South of the Antarctic Convergence in the Pacific Quad- rant of the Antarctic Ocean; Eltanin station 58-921; 5' Blake Trawl; 68°51'— 68°56’S; 114°08’—114°10’W; 4,038 m; 16 January 1964. Derivation of name.—The name is the Latin neuter adjective meaning antarctic and it modifies the neuter Mesosignum. Disposition of type.—The type-specimen has been deposited in the Na- tional Museum of Natural History (holotype female USNM 171447). Affinities. —The new species has long posterolateral extensions on the pleotelson and is thus in the Type B group of Menzies and Frankenberg (1968). Only M. ansatum Menzies and Frankenberg lacks short spines on the dorsum which is comparable to the very few short spines present on the dorsum of the new species. The posterolateral extensions on segments II and III are comparatively much shorter on M. antarcticum. On peraeonal segments IV, V and VI posterolateral extensions are not even suggested as they are in M. ansatum. The new species can be told from all others in the genus because it has only a few small spines on the dorsum and on the peraeonal extensions. This work was done under contract with the Smithsonian Oceanographic Sorting Center (PC-808755, Fund 16612600-P10000-256). Literature Cited Birstein, J. A. 1963a. Deep water isopods (Crustacea. Isopoda) of the north-western part of the Pacific Ocean.—Akad. Nauk. USSR Trudy Inst. Okeanolog. pp. 1-214. . 1963b. Isopods (Crustacea, Isopoda) from the ultra-abyssal zone of the Bougainville Trench.—Zool. Zhur. 46(6)814—834. Menzies, R. J. 1962. The isopods of abyssal depths in the Atlantic Ocean.—In: Abyssal Crustacea. *“‘Vema’’ Res. Ser., Columbia Univ. Press 1:80—206. and D. Frankenberg. 1968. Systematics and distribution of the bathyal-abyssal genus Mesosignum.—Biol. Antarctic Seas III, Antarctic Res. Ser. 11:113-140. , and R. Y. George. 1972. Scientific Results of the Southeast Pacific Expeditions.— Isopod Crustacea of the Peru-Chile Trench. ‘‘Anton Bruun”’ Rept. 9:1—124. 15 Smith St., Hampton, N.J. 08827. PROC. BIOL. SOC. WASH. 92(2), 1979, pp. 333-338 A NEW WATER SCAVENGER BEETLE FROM MEXICO (COLEOPTERA: HYDROPHILIDAE) Paul J. Spangler and Silvia Santiago-de Bueno Abstract.—A new hydrophilid beetle, Berosus gordito, new species, is described from the state of Durango, Mexico. Ten distinguishing morphlog- ical characters which serve to separate this new taxon from related species in the genus are described and figured. The biotopes are briefly discussed and two are illustrated. This new species of Berosus is described below to make the name avail- able for use in future studies and publications on water beetles of Mexico. The species is known, thus far, only from a few localities in the state of Durango near the city of Durango. Specimens originating from the collec- tions of the U.S. National Museum of Natural History are indicated by (USNM); those from the collections of the Instituto de Biologia, Universi- dad Nacional Autonoma de Mexico, are indicated by (UNAM). Berosus gordito, new species (Figs. 1-12) This new species may be distinguished by the following combination of characters: head black, pronotum and scutellum testaceous; mesosternal crest toothed posteriorly and sinuous behind tooth; female with 2 distinct teeth in apical emargination of fifth abdominal sternum and elytral apices prolonged into acute angles; male with 2 small teeth in emargination of fifth abdominal sternum but obscured by an apicomedial lobe; elytral apices nor- mal, not prolonged or angular; apices of male parameres each with a small but distinct transverse reflexed ridge dorsally at apex. Length of holotype male 4.5 mm, greatest width 3.0 mm. Color of head black, with brassy reflection. Pronotum testaceous. Elytron testaceous but apical and anterolateral areas slightly lighter; humeral macula light brown; interval 4 with faint brownish macula before midlength; intervals 3 and 4 each with a dark brown macula slightly behind midlength; intervals 4 and 6 each with a dark brown linear macula at apical four-fifths; elytral striae dark brown to piceous. Scutellum testaceous. Venter black except proster- num and procoxae ferrugineous; apex of mesosternum and mesocoxae pi- ceous; other segments of legs testaceous; tibial spurs and tarsal claws dark reddish brown. Maxillary palpus testaceous, except apex infuscate. Antenna and labial palpus testaceous. 334 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON € € € e 2 race pesoe rene G Sac eel ects © CLAS 0.5mm > @ ° ee ee el 2 Figs. 1-7. Berosus gordito, n. sp.: 1, emargination of fifth abdominal sternum, 6, vv; 2, same of 2; 3, tarsal claw, lv; 4, mesosternal crest, lv; 5, metasternal process, vv; 6, elytral apices, 6, dv; 7, same of 2. (vv = ventral view; lv = lateral view; dv = dorsal view). Head moderately coarsely punctate, punctures separated by one-half to one time their width; labrum with fine and dense seta-bearing punctures; setae more conspicuous apically. Pronotum with very feeble alutaceous sculpture on disc, becoming stronger laterally; subrectangular, slightly more than twice as wide as long; finely margined anteriorly and laterally, antero- lateral and posterolateral angles rounded; sides feebly arcuate; discal punc- tures finer than those on head; lateral punctures coarser, similar to those on head. Scutellum with punctures similar in size and density to lateral punctures on pronotum. Elytron with 11 striae, discal striae distinctly im- pressed, lateral striae feebly so, second stria incomplete, not attaining mid- length; striae 7 and 8 arising behind humeral macula, not from base; most punctures of striae and intervals slightly larger than discal punctures of pronotum; punctures of intervals fine and coarse intermixed; apex and margin with seta-bearing punctures; humeral angles smoothly rounded; sides finely VOLUME 92, NUMBER 2 335 10 Figs. 8-10. Berosus gordito, n. sp., male aedeagus: 8, basal piece, lv; 9, apical piece, lv; 10, apices of parameres and median lobe, dv. (lv = lateral view; dv = dorsal view). margined; apical angles rounded; sutural angle entire (Fig. 6). Legs pubes- cent on basal half of profemur and mesofemur; metafemur pubescent on basal three-fifths. Protarsus four segmented; basal segment subequal to sec- ond and third segments combined, moderately expanded, with pubescent pad along entire length ventrally; second segment short, subequal to third, 336 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON slightly expanded, with pubescent pad on apicolateral angle ventrally; third segment not expanded and without pubescent pad; fourth segment as long as second and third segments combined, without pubescence. Tarsal claws long, slender, each claw with a small acute basal tooth (Fig. 3). Venter with short, fine, dense, yellowish hydrofuge pubescence. Mesosternal crest (Fig. 4) low anteriorly, moderately high posteriorly, with moderately large pos- teroapical tooth, and sinuous behind tooth. Metasternal process (Fig. 5) with acute apicolateral projections shorter than posteromedial crest. Ab- dominal sterna not carinate; fifth abdominal sternum with two small median teeth obscured by an apicomedial lobe (Fig. 1). Aedeagus as illustrated (Figs. 8, 9, 10). Female.—The allotype differs from the male as follows: pronotum dis- tinctly alutaceous; protarsus five segmented, without pubescent pads; fifth abdominal sternum with two small median teeth in apical emargination (Fig. 2) and apicomedial lobe absent, therefore, teeth more distinct than those on male; elytral apices (Fig. 7) moderately produced, acutely angular, and over- lapping in normal position. Variations.—This taxon is variable in a number of characters as follows: Length from 4.5 to 6.0 mm; greatest width 3.0 to 4.1 mm. The brownish maculae on the elytra, as in many species of Berosus, vary widely from very reduced maculation to moderately confluent maculae; especially those on intervals 3, 4, and 6. When these maculae (3, 4, 6) are confluent, they become subvittate, i.e., the confluency is longitudinal, not transverse. In teneral and lightly pigmented specimens the humeral and anterior maculae on interval 3 (actually 2 and 3 combined) are often absent, and in these instances all maculae usually are restricted to the posterior half of the ely- tron. In lightly maculate specimens, interval 5 usually is immaculate; but in more obviously maculate forms, interval 5 may have a medial and rarely a second macula just before midlength. The posteriorly abbreviated second elytral strial row of punctures usually is straight and terminates on midline between first and third striae but in some examples this stria curves inward and terminates by merging with the first stria. The alutaceous pronotal sculp- ture is distinct on all females examined. On males this sculpture is much more feeble and in a few (3 out of 17) essentially absent. Type-data.—Holotype male: MEXICO: Durango, Cerro Gordo, 28 June 1964, Paul J. Spangler, USNM Type No. 75664, deposited in the National Museum of Natural History, Smithsonian Institution. Allotype, same data as holotype. Paratypes: MEXICO: Same data as holotype, 13¢¢, 12922 (USNM); Durango, Morcillo (at Lago Pena del Aguila), 28 June 1964, Paul J. Spangler, 16, 12 (USNM): Durango, Durango (25 KM west), 29 June 1964, Paul J. Spangler, 1¢ (USNM): Durango, Guatimapé, Laguna Santia- guillo, 18 August 1970, G. Fernandez 26 3,222 (UNAM). VOLUME 92, NUMBER 2 357, Fig. 11. Berosus gordito, n. sp., biotope, type-locality. Etymology.—The name gordito is from the Spanish vernacular meaning little fat one and is used here in apposition to the generic name. The name appropriately describes the shape of this species which is stouter than the majority of the species of Berosus when viewed from above. This new taxon is one of only 4 species of Berosus described from Mexico and Central America which have a testaceous pronotum and scutellum. Two of these species, B. stramineus Knisch and B. metalliceps Sharp, may be eliminated immediately by the lack of teeth in the emargination of the fifth abdominal sternum. Therefore, B. gordito is most similar to B. rubellus Knisch because both have 2 teeth in the emargination. Berosus gordito may be distinguished from rubellus as follows: mesosternal crest ascending to the tooth, then descending and sinuous behind tooth; elytral apices pro- duced and acutely angular in female; first abdominal sternum ecarinate; teeth in emarginate fifth sternum obscured by an apicomedial lobe and ely- tral apices normal in males. Conversely, rubellus has: a truncate mesoster- nal crest; male and female elytral apices normal; first abdominal sternum carinate and faintly extending to basal third; teeth in emargination fine but no apicomedial lobe obscuring them in males. Habitat.—The specimens of the type-series from Cerro Gordo were col- lected from shallow roadside pools adjacent to pastures (Fig. 11). These pools were enriched with cow manure and bordered with Marsilea and 338 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 12. Berosus gordito, n. sp., biotope. Lago Pena del Aguila. Jussiaea. The specimens from Morcillo (at Lago Pena del Aguilla) (Fig. 12) were collected from a pool at the base of the dam breast. The specimens from 25 miles west of Durango were collected in roadside ditches. Acknowledgments We extend our thanks to Mr. Michael Druckenbrod, Smithsonian Insti- tution staff artist for preparing the line drawings for this article. Thanks also are gratefully extended to the National Science Foundation for Grant GB- 1697 which enabled Spangler to collect this and other new species in Mexico and Central America. (PJS) Smithsonian Institution, Washington, D.C. 20560; (SS-deB) Insti- tuto de Biologia, UNAM, Aptdo. 70-153, Mexico 20, D.F. PROC. BIOL. SOC. WASH. 92(2), 1979, pp. 339-359 JUVENILE MORPHOLOGY OF THE RARE BURROWING MUD SHRIMP NAUSHONIA CRANGONOIDES KINGSLEY, WITH A REVIEW OF THE GENUS NAUSHONIA (DECAPODA: THALASSINIDEA: LAOMEDIIDAE) Joseph W. Goy and Anthony J. Provenzano, Jr. Abstract.—The status and history of the genus Naushonia is reviewed. Early juvenile stages of N. crangonoides reared in the laboratory from larvae captured in the plankton afforded an opportunity to evaluate changes in morphology with size. The fifth juvenile stage is described in detail and compared with adults. Some taxonomic characters used previously to dis- criminate species within this genus are invalid but others allow separation of the species. Comparisons of N. crangonoides with the two other Amer- ican species and with description of the Red Sea species permitted the construction of a key for the identification of the known species of the genus. Introduction The genus Naushonia was erected by Kingsley (1897) for a small male shrimp found by Professor Hermon C. Bumpus in the sand of the channels of the Island of Naushon off the coast of Massachusetts. Kingsley named this shrimp Naushonia crangonoides because of certain morphological sim- ilarities to the Crangonidae, but noted differences which might subsequently justify the establishment of a new family. A short note was published by Gray (1901) on a second specimen of this species, an ovigerous female, collected by himself in the sand of the shore of Ram Island, near Woods Hole, Massachusetts. Thompson (1903) redescribed these two specimens and also described some unusual late larval stages taken in the plankton off Woods Hole. Some of them metamorphosed in the laboratory allowing him to attribute these planktonic larvae to N. crangonoides. Thompson re- marked on the similarity of his zoeae to those of Calliaxis adriatica (=Jaxea nocturna) from the Mediterranean, and suggested placing Naushonia in the Family Laomediidae of the Thalassinidea. Chace (1939) synonymized with Naushonia the genus Homoriscus Rath- bun, containing two species, H. portoricensis (Rathbun, 1901) and H. mac- ginitei (Glassell, 1938) and included Coralliocrangon perrieri (Nobili, 1904) in Naushonia as well. Chace devised a tentative key to separate the four 340 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON species and stated the need for a re-examination of the Massachusetts (N. crangonoides) and Red Sea (N. perrieri) species. From the descriptions and illustrations of N. crangonoides by Kingsley (1897) and Thompson (1903), it is difficult to differentiate this type-species of the genus from the other three known species. In a previous paper (Goy and Provenzano, 1978), we redescribed the early larval development of N. crangonoides. During that study, we reared the fifth juvenile stage of N. crangonoides from captured planktonic first stage zoeae. We take this opportunity to present a description and illustrations of the juvenile morphology of N. crangonoides, to compare it with adult spec- imens and to summarize the differences between the four known species of the genus. Acknowledgments We are grateful to Dr. Raymond B. Manning, Curator, Department of Invertebrate Zoology, Smithsonian Institution, who made it possible for us to examine specimens of Naushonia crangonoides, N. macginitei, N. por- toricensis, the paratype of Homoriscus (=Naushonia) macginitei and the type of Homoriscus (=Naushonia) portoricensis. Dr. Austin Williams pro- vided us with a juvenile N. crangonoides collected from Bogue Sound, North Carolina. This work was supported by National Science Foundation Grant number DEB-76-11716. Methods and Material In our earlier paper, we mentioned that some individuals lived beyond the postlarval stage. Of these, two individuals survived to the fourth juvenile stage and one molted to the fifth juvenile stage. These animals provided basic material for the study of the very early juvenile morphology described herein. USNM refers to catalog numbers of the National Museum of Natural History, Smithsonian Institution, and UNC-IMS to catalog numbers of the University of North Carolina Institute of Marine Science. The description of the Naushonia crangonoides postlarva has been presented earlier (Goy and Provenzano, 1978). Juveniles and exuviae of known history were preserved in 70% ethyl alcohol. Dead animals were heated slowly in 5% KOH for approximately ten minutes to remove tissue from the exoskelton. These specimens and all casts from molted animals were stained in either Mallory’s Acid Fuchsin Red or Chlorazol Black E (1% in 70% Alcohol). Appendages were dissected in lactic acid and mounted in glycerin jelly. Drawings were made with the aid of a camera lucida; measurements were made with the aid of a stage micrometer. Carapace length (CL) was measured from tip of rostrum to the posterolateral margin of the carapace. Total length (TL) was measured from VOLUME 92, NUMBER 2 34] the tip of the rostrum to the most posterior margin of the telson, and ex- cluded all telson processes and setae. Juvenile Stages of Naushonia crangonoides In the first few molts after the postlarval stage, the morphology of Nau- shonia crangonoides does not change drastically from that of the postlarva. There is a gradual development of adult characteristics. The following are the major morphological changes that occur from the first juvenile stage through the fifth juvenile stage. The telson and uropods are unchanged until the fifth juvenile stage is reached. In the first juvenile stage, the antennule is similar to that of the postlarva, but the 6 aesthetascs are now located on the 3 middle segments of the external flagellum, 2 per segment. This appendage does not change further until the fifth juvenile stage. The antenna of the first juvenile stage is unchanged from the postlarval stage, except that there are 20 plumose setae on the inner margin of the scale. This appendage also does not change appreciably until the fifth ju- venile stage. The mandibles show the most significant changes with each succeeding molt. The postlarval stage has symmetrical mandibles (Fig. 1A) with the cutting edge provided with 4 small teeth. The palp is developed but unseg- mented, bearing a minute seta terminally. In the first juvenile stage, the cutting edge of the mandible (Fig. 1B) has the 4 teeth of the postlarva plus a small medial tooth, and the palp is still unsegmented but now bears 2 terminal rows of 5 spines. In the second juvenile stage, the mandible (Fig. 1C) has a cutting edge with 8 terminal and 2 subterminal teeth. The palp is now 2-segmented, with the first segment bearing an outer long plumose seta and the second segment having 13 terminal stout spines. The third juvenile stage has the mandible (Fig. 1D) basically unchanged from the preceding Stage, except the cutting edge has only 10 small terminal teeth and the second segment of the palp bears 14 terminal spines. In the fourth juvenile stage, the mandible (Fig. 1E) has 4 large and 9 smaller teeth on the cutting edge. The palp shows signs of a third segment but is still 2-segmented, with 2 long plumose setae on the outer margin of the first segment and 15 terminal spines on the second segment. The maxillule of the juvenile stages is unchanged from the postlarval stage, except more spines and plumose setae develop on the coxal and basal endites with each succeeding molt. The maxilla of the juvenile stages is also similar to that of the postlarva. The number of setae increases on the 4 inner lobes, the endopodite and the scaphognathite. The endopodite remains unsegmented and by the second 342 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 1. Naushonia crangonoides: Mandibles of postlarva (A); First juvenile (B); Second juvenile (C); Third juvenile (D); Fourth juvenile (E); and Fifth Juvenile (F). juvenile stage the long tapering proximal lobe of the scaphognathite has developed 5 long whip-like plumose setae. The first maxilliped shows some significant changes with each successive molt past the postlarva. In the first juvenile stage, the basipodite and en- dopodite remain unchanged from the postlarva. The expodite is still 3-seg- mented with the proximal segment now bearing 12 plumose setae and the terminal segment having 3 long plumose setae. The epipodite is large, serrate and triangular. In the second juvenile stage, the endites of the basipodite increase their numbers of setae and the endopodite is now 2-segmented with the distal segment enlarged, rounded and bearing 5 plumose setae. The exopodite is now 5-segmented with the terminal segment bearing only 2 long plumose setae. The first maxilliped does not show further change until the fifth juvenile stage. The second maxilliped shows a more gradual change to the adult appen- dage after the postlarval stage. In the first juvenile stage, the endopodite becomes 5-segmented but the rest of the appendage is unchanged from the preceding stage. By the second juvenile stage, the endopodite has the pen- ultimate segment expanded and the exopodite bears 5 long plumose setae 343 VOLUME 92, NUMBER 2 Sess SS, SAyad \ ~\ Fig. 2. Naushonia crangonoides: Fifth juvenile stage: Lateral view (A); Dorsal view (B): Antenna (C); and Antennule (D). 344 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON on the terminal segment. This maxilliped does not change considerably until the fifth juvenile stage. The third maxilliped is unchanged from the postlarval stage in the first juvenile stage, except there are now 2 long plumose whip-like setae on the basipodite and the ischium distally bears 5 teeth. There is also an increase in setae on the segments of the endopodite. In the second juvenile stage, the teeth of the serrate ischium increase to 10 but the remainder of the appendage is the same until the fifth juvenile stage. None of the pereiopods changes significantly after the postlarval stage, but all gradually become larger and more setose. The pleopods remain unchanged from the postlarva until the fifth juvenile Stage. Fifth Juvenile Stage The one animal that molted to this stage (TL 6.0 mm, CL 2.3 mm) died after 15 days. Carapace (Figs. 2A, 2B) cylindrical, depressed in front with rostrum slightly down curved. Rostrum triangular, flat, extending beyond eyes with finely serrate recurved borders. Anterior borders of carapace serrate with supraorbital and antennal spines. Dorsal and branchial areas distinguished by straight, prominent, longitudinal groove (linea thalassinica) and cervical groove also well marked in middle. Carapace smooth except along grooves and ridges, with posterior margin bearing 20 fine hairs. Eyes still visible from above with minute pigment spot. Abdomen about a third longer than carapace, smooth without spines or carinae. Borders of pleura of first and sixth segments truncate, those of other segments rounded. Peduncle of antennule (Fig. 2D) composed of 4 segments extending be- yond front of eyes. Proximal segment with 14 long plumose setae on outer margin and ring of 12 setae subterminally on inner edge. Second segment ends in 2 spines and bears 5 feathered setae on outer margin, 1 medially and 1 terminally. Third segment ends in spine on inner border with medial spine and bears 5 plumose terminal setae. Fourth segment with 2 medial and 3 terminal plumose setae. External flagellum of 5 segments, with first segment lacking setae or aesthetascs. Next 3 segments each bear 2 aesthetascs on outer margin and 4, 1 and 3 setae respectively. Terminal segment with 3 plumose setae at apex. Inner flagellum 4-segmented with 2 setae on each segment, but terminal segment bearing 4 setae. Peduncle of antenna (Fig. 2C) with 5 segments. First segment with 2 spines on outer margin, 1 spine on inner margin, and bearing antennal scale. Next 2 segments with spine on each margin while penultimate and ultimate segments lack spines or setae. Antennal scale ovate with 5 outer teeth, 7 VOLUME 92, NUMBER 2 345 Fig. 3. Naushonia crangonoides: Fifth juvenile stage appendages: Maxilla (A); Maxillule (B); and First maxilliped (C). 346 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON \ a ANG 4; / PRD VWI, See EY SS > SSS = ae eS <=> p> es Fig. 4. Naushonia crangonoides: Fifth juvenile stage appendages: Telson (A); Second max- illiped (B); and Third maxilliped (C). VOLUME 92, NUMBER 2 347 outer setae and 17 plumose setae on inner margin. Flagellum long and ta- pering consisting of 51 segments, most having circlet of setae at distal ends. Mandible (Fig. 1F) with only 2 large and 6 smaller teeth on cutting edge. Palp 3-segmented, with second segment having 3 long plumose setae and terminal segment bearing 24 spines. Maxillule (Fig. 3B) with 2-segmented endopodite with flexed terminal seg- ment and with first segment bearing plumose seta terminally and one on inner margin. Basipodite bearing 4 teeth and 2 plumose setae on border. Coxal endite with 30 feathered setae marginally and submarginally. Basal endite with 30 feathered setae marginally and submarginally. Basal endite rounded with 2 and 4 plumose setae on outer and inner margins, respec- tively; terminally 19 stout teeth and 3 long plumose setae, and subterminally 9 shorter plumose setae. Maxilla (Fig. 3A) with following setation on 4 inner lobes: 11 on proximal lobe of coxal endite; 8 on distal lobe; 15 on proximal lobe of basal endite; 21 on distal lobe. Unsegmented endopodite bearing 4 plumose setae distally and 2 proximally on outer margin, and 2 plumose setae on inner margin. Scaphognathite broad with 60 plumose setae on outer margin; also 22 minute setae on inner margin. Long tapering proximal lobe bearing 4 long plumose setae and 3 longer, whip-like plumose setae. First maxilliped (Fig. 3C) with 2-lobed basipodite; proximal lobe with 20 plumose setae and 21 shorter non-plumose setae; distal lobe bearing 52 plumose setae. Endopodite 2-segmented with distal segment expanded and triangular, bearing 5 long plumose setae and 8 shorter plumose setae. Ex- opodite 7-segmented with proximal segment widened and bearing 15 long feathered setae on outer margin. Last 5 segments each with long plumose seta on both sides near base, except terminal segment also with 4 long plumose setae at apex. Epipodite large, serrate and triangular with arthro- branch and mastigobranch. Second maxilliped (Fig. 4B) with 5-segmented endopodite with penulti- mate segment slightly expanded. These segments bearing long plumose se- tae proximally to distally as follows: 4; 24; 2; 10 and 16. Exopodite 7-seg- mented with first segment having numerous short plumose setae and last 5 segments bearing long plumose setae; terminal segment having 4 setae at apex. Epipodite heavily serrate, with 2 arthrobranchs and podobranch. Third maxilliped (Fig. 4C) having endopodite of 5 segments with ischium bearing 13 prominent teeth on inner border and numerous plumose setae. Second segment with 3 prominent teeth on outer border besides numerous plumose setae; other 3 segments heavily setose. Exopodite consisting of 7 segments with last 4 segments bearing long plumose setae. Epipodite com- plex consisting of small anterior lobe and bunch of long, plumose whip-like setae, serrate-margined mastigobranch and podobranch. Also 2 arthro- branchs on third maxilliped. 348 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Telson (Fig. 4A) with rounded end, no longer possessing spine at each external angle as in postlarval and preceding juvenile stages. Outer margin bearing 40 plumose setae with numerous submarginal and medial shorter setae. Uropods with serrate transverse sutures on both rami that end with external spine. Endopodite and exopodite both with 40 plumose setae on outer borders. Exopodite also with six plumose setae submarginally and five external spines ending in a stout movable spine. Chelipeds (Fig. 5A) large, slender, subchelate. Ischium slightly smaller than merus with 5 small teeth on inner border. Merus with 2 small outer teeth and 4 small inner teeth plus large spine near apex. Carpus triangular in outline, articulating with propodus by 2 tubercles. Propodus setose, elon- gate, bearing 3 prominent teeth and 6 smaller teeth on distal inner margin, 2 small teeth terminally, and 13 small teeth along entire length of outer border. Dactylus bent at base almost at right angle, very slender and falcate, and with sharp margins, outer of which fringed with long setae. Two ar- throbranchs, small podobranch, and slender mastigobranch present. Second pereiopods (Fig. 5B) short, flattened and setose on ventral margin. Dactylus robust, bearing 10 small teeth on inner margin and numerous long setae on outer margin. Third pereiopods (Fig. SC) longer than fourth and fifth legs (Figs. 5D, SE) but all 3 pairs slender with long propodi and arcuate dactyli; those of third pair bearing 18 small teeth on inner margins. Two arthrobranchs on second, third and fourth pairs of pereiopods, small podobranch on second and third pairs, and slender mastigobranch on all 3 pairs. No gills on fifth pereiopods. Pleopods (Figs. 5F, 5G, 5H, 51) absent on first abdominal somite but present on second to fifth somites. Pleopods biramous, lanceolate and with- out stylambys. Endopodites and exopodites with 12 and 14 long plumose setae respectively. Museum Specimens Naushonia crangonoides 1.—CL 7.5 mm; TL 21.0 mm. (USNM 34143). Male. Eyes barely visible from above. No spines on telson. Uropods with complete transverse su- tures, exopod with 5 spines on lateral margin ending in a strong movable spine. Antennal scale margin with 11 teeth. Mandibular palp 2-segmented. Third pereiopod has 12 and fourth pereiopod has 7-10 movable spines on outer margins of dactyli. 2.—CL 6.5 mm; TL 17.5 mm. (USNM 102277). Female. Eyes visible from above. Telson, uropods, antennal scale and mandibular palp same as above. Third pereiopod has 20 and fourth pereiopod has 12 movable spines on dactyli. 3.—CL 10.0 mm; TL 25.0 mm. (USNM 102279). Female. Eyes not visible VOLUME 92, NUMBER 2 349 Fig. 5. Naushonia crangonoides: Fifth juvenile stage appendages: Cheliped (A); Second pereiopod (B); Third pereiopod (C); Fourth pereiopod (D); Fifth pereiopod (E); Second pleopod (F); Third pleopod (G); Fouth pleopod (H); and Fifth pleopod (I). 350 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON from above. Telson, uropods, antennal scale and mandibular palp same as above. Third pereiopod has 24 and fourth pereiopod has 15 movable spines on dactyli. 4.—CL 11.0 mm; TL 27.0 mm. (USNM 102280). Female. Eyes not visible from above. Telson, uropods, antennal scale and mandibular palp same as above. Third pereiopod has 20 and fourth pereiopod has 14 movable spines on dactyli. 5.—CL 4.5 mm; TL 11.0 mm. (UNC-IMS 254). Female. Eyes visible from above. Telson and uropods same as above. Antennal scale margin with 7 teeth. Mandibular palp 3-segmented. All pereiopods missing. Naushonia portoricensis 1.—CL 6.3 mm; TL 14.5 mm. (USNM 23782). Female. Type of Homoris- cus (=Naushonia) portoricensis. Eyes visible from above. Telson with spine on lateral margin. Uropods with complete sutures, exopod with 2 spines on lateral margin ending in a strong movable spine. Antennal scale margin with 6 teeth, distal tooth largest and curved inward. Mandibular palp 3-segmented. All pereiopods missing. 2.—CL 4.2 mm; TL 12.0 mm. (USNM 155101). Male. Eyes visible from above. Telson, uropods, antennal scale and mandibular palp same as above. Third pereiopod has none and fourth pereiopod has 20 movable spines on dactyli. 3.—CL 2.0 mm; TL 5.6 mm. (USNM 155101). Male. Eyes visible from above. No spines on telson. Uropods and mandibular palp same as above. Antennal scale margin with only 4 teeth. All pereiopods missing. Naushonia macginitiei 1.—CL 7.6 mm; TL 19.0 mm. (USNM 171605). Ovigerous female. Para- type of Homoriscus (=Naushonia) macginitei. Eyes visible from above. Telson with 3 spines on lateral margin. Uropods with incomplete transverse sutures, exopod with 2 spines on lateral margin, ending with 2 small spines and a large movable spine. Antennal scale margin with 7 teeth. Mandibular palp 3-segmented. Third pereiopod has 20—22 and fourth pereiopod has 18 movable spines on dactyli. 2.—CL 7.5 mm; TL 18.2 mm. (USNM 171604). Female. Eyes visible from above. Telson, uropods and mandibular palp same as above. Antennal scale margin with 8 teeth. Third pereiopod has 20-24 and fourth pereiopod has 16 movable spines on dactyli. 3.—CL 7.5 mm; TL 19.1 mm. (USNM 144492). Female. Eyes visible from above. Telson, uropods and antennal scale same as paratype. Third pereio- pod has 22 movable spines on dactyli. Fourth pereiopod missing. 4.—CL 6.5 mm; TL 17.2 mm. (USNM 144492). Female. Eyes visible from above. Telson, uropods and mandibular palp same as paratype. Antennal VOLUME 92, NUMBER 2 351 scale margin with 8 teeth. Third pereiopod has 24 and fourth pereiopod has 15 movable spines on dactyli. Systematic Position of Naushonia Order Decapoda Supersection Macrura Reptantia Section Thalassinidea The Decapod section Thalassinidea is considered a valid group but there is much difference of opinion as to its number of families and its systematic position. Balss (1957) recognized only 4 families, but earlier Gurney (1938) recognized 7 distinct families based on a combination of adult and larval characteristics: the Axianassidae, Axiidae, Callianassidae, Callianideidae, Laomediidae, Thalassinidae and Upogebiidae. Gurney discussed these fam- ilies in detail and gave a key to their determination. The thalassinids are all burrowing forms characterized by a well calcified, compressed carapace; a symmetrical, extended, often feebly calcified abdomen ending in a well de- veloped tail fan; first pereiopods chelate or subchelate, second chelate, sub- chelate or simple, and third legs always non-chelate (Wear and Yaldwyn, 1966). Family LAOMEDIIDAE Borradaile, 1903 The family Laomediidae at present consists of 3 thalassinid genera and 8 described species. According to Wear and Yaldwyn (1966), these are char- acterized by ‘‘having a linea thalassinica; first legs subequal and chelate or subchelate; second pereiopods subchelate or simple; no appendix interna on pleopods; uropods with transverse sutures; podobranchs on at least sec- ond and third maxillipeds and first and second pereiopods, and epipods on the first to fourth pereiopods.’’ Chace (1939) divides the Laomediidae into 2 subfamilies: Laomediinae and Naushoniinae. The Naushoniinae are dis- tinguished from the Laomediinae by the subchelate rather than chelate first legs, the well-developed antennal scale that is absent or rudimentary in the Laomediinae, and the simple instead of subchelate last pereiopods. De Man (1928) considered the family Axianassidae untenable and included its only genus (Axianassa) in the family Laomediidae. However, the two species of this genus (Axianassa intermedia and A. mineri) lack some of the major characters of the laomediids, including transverse sutures on the uro- pods and exopodites on the third maxillipeds. According to Yaldwyn and Wear (1972), all examined adult species of the family Laomediidae have 18 gills though there may be some minor differences in their distribution. Both species of Axianassa have only 17 gills. Therefore, in the present study, only Jaxea, Laomedia, and Naushonia are included in the Laomediidae. 352 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Subfamily LAOMEDIINAE Jaxea Nardo, 1847 is known from 2 named species, J. nocturna from the Mediterranean and North Atlantic (deMan, 1928; Zariquey-Alvarez, 1968) and J. novazealandiae from New Zealand (Wear and Yaldwyn, 1966). Laomedia de Haan, 1849 is known from 2 described species, L. astacina from Korea and Japan (deMan, 1928; Sakai, 1962) and L. healyi from eastern Australia (Yaldwyn and Wear, 1972). A third species of Laomedia, still undescribed, was found in eastern Australia by Yaldwyn and Wear (1972) during their study of L. healyi. Larvae from Sydney Harbour attrib- uted to Jaxea sp. by Dakin and Colefax (1940) probably belong to one of these eastern Australian species of Laomedia. A first stage larva from Sa- moa believed by Gurney (1938) to belong to Jaxea sp. also may represent an undescribed species of Laomedia since this larva has characters more similar to Laomedia than to Jaxea (Sakai and Miyake, 1964; Goy and Pro- venzano, 1978). Subfamily NAUSHONIINAE The genus Naushonia Kingsley, 1897 is Known from 4 species: N. cran- gonoides from off Massachusetts (Kingsley, 1897); N. portoricensis from Puerto Rico (Rathbun, 1901); N. perrieri from the Red Sea (Nobili, 1904); and N. macginitiei from southern California (Glassell, 1939). Larval stages probably belonging to N. portoricensis were described by Gurney and Le- bour (1939) from Bermuda and larvae found off Samoa and the Great Barrier Reef (Gurney, 1938) might belong to N. perrieri. There are at least 2 addi- tional, apparently separate, species of Naushonia that are undescribed and known only from their larvae, off New South Wales (Dakin and Colefax, 1940) and from the Adriatic Sea (Kurian, 1956; Goy and Provenzano, 1978). Review of the Genus Naushonia Naushonia was founded by Kingsley (1897) for a single adult male spec- imen collected in the sand on Naushon Island, near Woods Hole, Massa- chusetts. A second adult, an ovigerous female, was collected by Gray (1901) from a 10 inch deep burrow in the sand on Ram Island, in Great Harbor, Woods Hole. Both of these specimens are now in the Gray Museum, Marine Biological Laboratory, Woods Hole. Four more adults are known from Massachusetts at Bass River, Vineyard Sound, and Elizabeth Islands (Wil- liams, 1974), which are in the USNM collection. Larvae believed to belong to N. crangonoides have been collected from the Woods Hole area during July, August, and September (Thompson, 1903; Fish, 1925); in Delaware Bay from August to October (Deevey, 1960); in Narragansett Bay in August (Hillman, 1964); and in Chesapeake Bay from August to September (San- VOLUME 92, NUMBER 2 353 Table 1. Comparison of some characters of first juvenile stages of Naushonia crangonoides and Naushonia portoricensis. N. crangonoides N. portoricensis Total Length 4.80 mm 3.25 mm Rostrum No lateral teeth Lateral tooth on side Rostral apical process Absent Present Linea thalassinica Distinct Faint Antennal scale 6 marginal teeth 3 marginal teeth Mandibular palp Unsegmented Three-segmented difer, 1972; Goy, 1976). In these last 2 collections, first stage larvae of Naushonia were most numerous near the bay mouth, suggesting a breeding population of N. crangonoides somewhere near the mouth of Chesapeake Bay. This theory was recently confirmed by Langton and Brodeur (1978) when they found 3 adult N. crangonoides in the stomach of a stingray, Dasyatis centroura, collected approximately 15 nautical miles northeast of the mouth of Chesapeake Bay. A juvenile N. crangonoides (TL 11.0 mm) has been collected in Bogue Sound, North Carolina (Williams, personal communication, 1977) extending the species known range over 700 nautical miles south of its type-locality. Chace (1939) synonymized with Naushonia the genera Homoriscus Rath- bun, 1901 and Coralliocrangon Nobili, 1904. This added 3 more species to Naushonia: N. portoricensis, N. macginitei, and N. perrieri. Naushonia portoricensis 1s known from Puerto Rico (Rathbun, 1901), Cuba (Chace, 1939), Bermuda (Gurney and Lebour, 1939) and is extended to Ascension Bay, Quintana Roo, Mexico from material examined in the present study. N. macginitei is known from southern California (Glassell, 1938) and is extended south to Ensenada de San Francisco, Sonora, Mexico from ma- terial examined in the present study. N. perrieri is known only from 2 specimens collected in French Somaliland, Red Sea. Chace (1939) devised a tentative key to the 4 species of Naushonia. From the results of the present study, this key can no longer be considered valid. There are lateral movable spines on at least the fourth pereiopod dactyl of all species. In N. crangonoides specimens less than 20 mm TL have their eyes visible from above. The postlarva to the fourth juvenile stage of N. crangonoides has the telson armed with a single pair of lateral spines like N. portoricensis. Also the fifth juvenile stage of N. crangonoides has a 3- segmented mandibular palp and an antennal scale with 5 marginal teeth, the same number as WN. portoricensis of similar size. There is a strong possibility that the ranges of N. crangonoides and N. portoricensis may overlap along the southeast coast of the United States. 354 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 6. Carapace of Naushonia crangonoides (A); N. portoricensis (B); N. macginitei (C); and N. perrieri (D). Mandibles of N. crangonoides (E); N. portoricensis (F); N. macginitei (G); and N. perrieri (H). (D and H adapted from Nobili, 1906.) VOLUME 92, NUMBER 2 355 Fig. 7. Chelipeds of Naushonia crangonoides (A); N. portoricensis (B; adapted from Chace, 1939); N. macginitei (C); and N. perrieri (D; adapted from Nobili, 1906). The early juvenile stages of N. crangonoides bear a close resemblance to specimens of the much smaller species N. portoricensis. The first juvenile stage of N. portoricensis described by Gurney and Lebour (1939) is quite similar to the first juvenile stage of N. crangonoides, but they show some differences (Table 1). Many of the characters given by Chace (1939) to separate these 2 species will overlap at the smaller size ranges. The antennal scale, telson and uropods at these sizes will enable one to differentiate the 2 species. The smallest N. portoricensis we examined (5.6 mm TL) has an antennal scale with 4 marginal teeth with the distal tooth curved inward, while the fifth juvenile stage of N. crangonoides (6.0 mm TL) has an an- tennal scale with 5 marginal teeth. The telsons of both these specimens have no external spines but the uropodal exopodite of N. portoricensis has 2 spines on the lateral margin which ends in a strong movable spine. The uropodal exopodite of the fifth juvenile stage of N. crangonoides has 5 external spines on the lateral margin which ends in a stout movable spine. The adults of the 4 species of Naushonia are generally similar in mor- phology but show differences in detail. Their carapaces (Figs. 6A, B, C, D) 356 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 8. Telsons of Naushonia crangonoides (A); N. portoricensis (B), and N. macginitei (C). Antennal scales of N. crangonoides (D); N. portoricensis (E); and N. macginitei (F). Dactyli of third and fourth pereiopods of N. crangonoides (G, H); N. portoricensis (1, J); N. macginitei (K, L); and N. perrieri (M; adapted from Nobili, 1906). are very much the same but that of N. macginitei is more granulose in the rostral area. The mandibles (Figs. 6E, F, G, H) are essentially the same, except that N. crangonoides has a fusion of segments in the palp giving the general appearance of a 2-segmented palp whereas the other species clearly have a 3-segmented palp. The chelipeds (Figs. 7A, B, C, D) show differ- ences. The ischium is toothed in N. portoricensis and N. macginitei but smooth in N. crangonoides and N. perrieri. N. crangonoides has teeth on the inner border of the merus, as does N. portoricensis and N. macginitei, but the inner border of N. perrieri’s merus is smooth. The outer margin of the merus is toothed in N. portoricensis, smooth in N. crangonoides and N. perrieri, and with 2 teeth at the base in N. macginitei. The propodus is very similar in N. crangonoides and N. perrieri but shows some differences in the other 2 species. In N. portoricensis, the inner border of the propodus is toothed below the prominent tooth and has its outer margin toothed along its entire length. In N. macginitei, the inner border of the propodus is smooth below the prominent tooth and the outer margin is only toothed on its upper length. The telson, uropods and antennal scales of the species of Naushonia seem to show the largest differences in morphology. In N. cran- VOLUME 92, NUMBER 2 a5i] gonoides, the telson (Fig. 8A) has no spines, the uropods have complete transverse sutures, and the uropodal exopodite has 5 spines on the lateral margin which ends in a strong movable spine. In N. portoricensis, the telson (Fig. 8B) has a spine on its lateral margin, the uropods have complete trans- verse sutures, and the exopodite of the uropod has 2 spines on its lateral margin which ends in a strong movable spine. The telson of N. macginitei (Fig. 8C) has 3 spines on its lateral margin, the uropods have incomplete transverse sutures, and the uropodal exopodite has 2 spines on its lateral margin ending with 2 small spines and a large movable spine. The antennal scale of N. crangonoides (Fig. 8D) bears 11 marginal teeth; the scale of N. macginitei (Fig. 8F) bears 7-8 marginal teeth; and the scale of N. portori- censis (Fig. 8E) bears 6 marginal teeth, with the distal tooth the largest and curved inward. Nobili (1904, 1906) did not adequately describe or illustrate the telson, uropods and antennal scale of N. perrieri to show differences from the other 3 species of Naushonia. Some or all of the dactyli of the last 3 pereiopods in Naushonia have lateral movable spines. These movable Spines are on the third and fourth pereiopods (Figs. 8H, G, K, L) of N. crangonoides and N. macginitei; on the fourth pereiopod (Fig. 8J) of N. portoricensis; and on the last three pairs of periopods (Fig. 8M) of N. per- rieri. Although more material of the Red Sea species needs to be examined, the following key will distinguish the 4 species: 1. Uropods with complete transverse sutures. A. Linea thalassinica pronounced, carinae of the carapace weak; telson without lateral spine; antennal scale with 10 or more mar- CUNIAIEUCCENE Me ee oe ee re re ees N. crangonoides B. Linea thalassinica not pronounced, carinae of carapace well marked; antennal scale with less than 10 marginal teeth. a. Telson with lateral spine; antennal scale with 6 marginal teeth, distal tooth largest and curved inwards; lateral movable spines on fourth pereiopod only .............. N. portoricensis b. Lateral movable spines present on all 3 posterior pereiopods Rar aM re Ie Bi eR NOT | Lehn Ae Sag 08 Ate eB ERE CES a! SR oe N. perrieri 2. Uropods with incomplete transverse sutures; uropodal exopodites with 2 lateral spines, margin ending with 2 small spines and stout MEOVADICSSPURE TR Me aioe aoe Ree eT ee ee eae N. macginitei Literature Cited Balss, J. 1957. Decapoda.—In: Bronn, H. G., Klassen und Ordnungen des Tierreichs, vol. 5, sec. 1, book 7, part 12:1505—1672. Borradaile, L. A. 1903. On the classification of the Thalassinidea.—Ann. Mag. Nat. Hist. ser. 7, 12:534—551. 358 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Chace, F. A., Jr. 1939. On the systematic status of the crustacean genera Naushonia, Ho- moriscus, and Coralliocrangon.—Ann. Mag. Nat. Hist. ser. 11, 3:524—530. Dakin, W. J. and A. N. Colefax. 1940. The plankton of the Australian coastal waters off New South Wales. Part IJ.—Publ. Univ. Sydney Dept. Zool. 1:1—125. Deevey, G. D. 1960. The zooplankton of the surface waters of the Delaware Bay region. Bull. Bingham Oceanogr.—Coll. Yale Univ. 17:5-53. Fish, C. J. 1925. Seasonal distribution of the plankton of the Woods Hole region.—Bull. U.S. Bur. Fish. 41:91-179. Glassell, S. A. 1938. New and obscure decapod Crustacea from the West American Coasts.— Trans. San Diego Soc. Nat. Hist. 8(33):411-454. Goy, J. W. 1976. Seasonal distribution and the retention of some decapod crustacean larvae within the Chesapeake Bay, Virginia.—Master’s Thesis, Old Dominion University, Nor- folk, Virginia, 334 pp. , and A. J. Provenzano, Jr. 1978. Larval development of the rare burrowing mud shrimp Naushonia crangonoides Kingsley (Decapoda: Thalassinidea; Laomediidae).—Biol. Bull. 154(2):241-261. Gray, G. M. 1901. Biological Notes.—Bull. U.S. Fish Comm. 19:307. Gurney, R. 1938. Larvae of decapod Crustacea. Part V. Nephropsidea and Thalassinidea.— Discovery Rep. 27:291-344. , and M. V. Lebour. 1939. The larvae of the decapod genus Naushonia.—Ann. Mag. Nat. Hist. ser. 11, 3:609-614. Hillman, N. S. 1964. Studies on the distribution and abundance of decapod larvae in Narra- gansett Bay, Rhode Island, with consideration of morphology and mortality.—Master’s Thesis, University of Rhode Island, Kingston, Rhode Island, 74 pp. Kingsley, J. S. 1897. On a new genus and two new species of macrurous Crustacea.—Bull. Essex Inst. 27:95—99. Kurian, C. V. 1956. Larvae of decapod Crustacea from the Adriatic Sea.—Acta Adriatica 6(3): 1-108. Langton, R. W., and R. D. Brodeur. 1978. Occurrence of a rare adult mud shrimp, Naushonia crangonoides Kingsley, 1897, in the Chesapeake Bay region (Decapoda, Thalassini- dea).—Crustaceana 35(1):96—98. deMan, J. G. 1928. The Thalassinidae and Callianassidae collected by the Siboga Expedition with some remarks on the Laomediidae.—Siboga Exped. Monogr. 39a6:1—187. Nobili, G. 1904. Diagnoses préliminaires de vingt-huit especes nouvelles de Stomatopodes et Décapodes Macroures de la Mer Rouge. Bull. Mus. d’Hist. Nat. Paris 10(5):228—238. . 1906. Faune Carcinologique de la Mer Rouge: Décapodes et Stomatopodes.—Ann. Sci. Nat. Zool. ser. 9, 4(1-3):82-85. Rathbun, M. J. 1901. The Brachyura and Macrura of Porto Rico.—Bull. U.S. Fish. Comm. for 1900, 2:1-127. Sakai, K. 1962. Systematic Studies on Thalassinidea. I. Laomedia astacina deHaan.—Publ. Seto Mar. Biol. Lab. 10(1):27-34. , and S. Miyake. 1964. Description of the first zoea of Laomedia astacina deHaan (Decapoda, Crustacea).—Sci. Bull. Fac. Agric. Kyushu Univ. 21(1):83-87. Sandifer, P. A. 1972. Morphology and ecology of Chesapeake Bay decapod crustacean lar- vae.—Ph.D. dissertation, University of Virginia, Charlottesville, Virginia, 532 pp. Thompson, M. T. 1903. A rare thalassinid and its larva.—Proc. Boston Soc. Nat. Hist. 31: 1—21. Wear, R. G. and J. C. Yaldwyn. 1966. Studies on thalassinid Crustacea (Decapoda, Macrura, Reptantia) with a description of a new Jaxea from New Zealand and an account of its larval development.—Zool. Publ. Victoria Univ. of Wellington 41:1—27. VOLUME 92, NUMBER 2 359 Williams, A. B. 1974. Marine flora and fauna of the northeastern United States. Crustacea: Decapoda.—Nat. Oceanic Atmos. Admin. Rep. Nat. Mar. Fish. Serv. Circ. 389, 50 pp. Yaldwyn, J. C., and R. G. Wear. 1972. The eastern Australian burrowing mud shrimp Lao- media healyi (Crustacea, Macrura, Reptantia, Laomediidae) with notes on the larvae of the genus Laomedia.—Aust. Zool. 17:126—141. Zariquey-Alvarez, R. 1968. Crustaceos Decapodos Ibericos.—Instit. Invest. Pesq. Pas. Nac. s/n Barcelona, 510 pp. (JWG) Duke University Marine Laboratory, Beaufort, North Carolina 28516; (AJP) Institute of Oceanography, Old Dominion University, Norfolk, Virginia 23508. PROC. BIOL. SOC. WASH. 92(2), 1979, pp. 360-367 OCTOPUS RAPANUI, NEW SPECIES, FROM EASTER ISLAND (CEPHALOPODA: OCTOPODA) Gilbert L. Voss Abstract.—Octopus rapanui, new species, is described from Easter Is- land. It is distinguished from other Indo-Pacific species by a combination of characters including the shape of the lower mandibular rostrum, long rachidian radular teeth, large secondary diverticulum of the penis and long, slender posterior salivary glands. In 1977 I received a small octopus for identification, collected at Easter Island by B. Alarcon and sent to me by Harald A. Rehder, Smithsonian Institution. The specimen was immature and in poor condition, but the beaks and radula were different from others I have examined from the Pa- cific. A search through the collections at the Rosenstiel School of Marine and Atmospheric Science, University of Miami, brought to light eight more specimens, some sexually mature but all in poor condition. These specimens were collected by Ian E. Efford and Jack Mathias, University of British Columbia, on the Medical Expedition to Easter Island. Examination of all of the specimens convinced me that they represent a new and undescribed species. A survey of the cephalopod literature has revealed no record of an octopod from Easter Island although the islands were visited by several of the major expeditions of the 18th and 19th centuries. Thus this species represents the first octopod to be included in the faunal reports of Easter Island. Octopus rapanui, new species Figs. 1, 2 Holotype.—Male, mantle length 115 mm, F82, rotenone at Hanga Piko, Easter Island, Efford and Mathias, leg., 14 January 1965, USNM 729860. Paratypes.—2 males, mantle lengths 97-113 mm, 1 female, mantle length 69 mm, F82, rotenone at Hanga Piko, Easter Island, Efford and Mathias, leg., 14 January 1965, USNM 729990.—1 female, mantle length 90 mm, M12, Apina Iti, Easter Island, Efford and Mathias, leg., 18 January 1965, UMML 1746.—1 male, mantle length 70 mm, 1 female, mantle length 107 mm, ‘‘collected in about 12 feet of water. Bottom rocky; in a sheltered cove. Caught at about 4:30 p.m.’’ Vinapu, Easter Island, Efford and Mathias, leg., 25 January 1965, UMML 1742.—1 male, mantle length 88 mm, rotenone in subtidal pool near camp at Hanga Roa, Easter Island, Efford and Mathias VOLUME 92, NUMBER 2 361 leg., S February 1965, UMML 1743.—1 female (?), mantle length 27 mm, Sta. E-37, in tide pool, B. Alarcon leg., Hanga Piko, Easter Island, 2 No- vember 1974, USNM 751587. Description.—The mantle is somewhat cylindrical and rather narrow ex- cept in one specimen (MWI males 41-49.8—74; females 51-—56.8—63); the head is set off from the mantle by a slight constriction. The head is narrow (HWI males 29-37.6—49; females 38-44.8-52) and bears small eyes. The mantle aperture is wide. The funnel is large, free for about half of its length. The funnel organ is indistinct but appears to be W shaped with the outer limbs a little longer than the median ones. The arms are long (MAI males 13-23.4-27; females 22—24.0-28: ALI males 75-80.6—96; females 76—78.5-80), stout basally but tapering to long slender tips. The arm order is usually I.IIJ.III.IV, but is somewhat variable, although I is almost always the longest and IV the shortest. In the females I is always the longest but in the males I was longest in two, II longest in three. The suckers are biserial and moderately large (SIn males 9-/0.2-11; females 9-//.3—15). There are no specially enlarged suckers in the males. The third right arm of the male is hectocotylized. It is shorter than the third left arm (HAI 72-84-95) and has a conspicuous spermatophoral groove. The ligula is small (LLI 1.4—2.8—4.0), medially excavated with thick- ened margins. The calamus is small but distinct (CLI 28.5-32./-—33.3). An interesting and possibly distinctive feature is best shown in the holotype. The spermatophoral groove at the base of the calamus widens and deepens into a pocket-like depression. It is visible but less distinct in other males of the species. The male with a mantle length of 88 mm had lost the major part of its hectocotylized arm but a new ligula had been produced that measured 1.8 mm complete with a calamus. The web order cannot be ascertained if there is one, for it is extremely variable. The web is rather low but it extends up the ventral side of the arms to near the distal third or further (WDI males 19—2/.2—22; females 18— 19.8—22). The gills bear 11 to 12 lamellae on the outer demibranch. The digestive tract was dissected out. It shows many unusual features. The buccal mass is more elongate than in the typical Octopus. The upper beaks are normal but the lower beaks have a nearly straight rostrum that projects forward like a spine. The radula is normal except for the very long and slender rachidian teeth. The rachidian shows a B,_- asymmetry. In the smallest specimen of 27 mm mantle length the rachidians are very long and slender with a possibility of three cusps on one side and two on the other. In the male of 88 mm mantle length the rachidians are somewhat shorter and broader with two to three cusps on one side with a B, order, while 362 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 1. Octopus rapanui: a, Lateral view of male, mantle length 88 mm; b, Radula of female, mantle length 107 mm; c, Radula of juvenile female (?), mantle length 27 mm; d, Digestive tract from female, mantle length 90 mm; e, Genitalia from male, holotype; f, Genitalia from female, mantle length 90 mm. VOLUME 92, NUMBER 2 363 Fig. 2. Octopus rapanui: a—b, Upper and lower mandibles of female, mantle length 107 mm; c—d, Upper and lower mandibles of male, mantle length 70 mm; e, Hectocotylus of holotype. there are only two cusps on the other side with a B,; order. In both specimens the third laterals are hooked at the tip. Both have marginal plates, those of the smaller poorly formed and indistinct. The anterior salivary glands are large, somewhat free from the buccal mass and closely embrace the esophagus. In the 88 mm length male these glands are darkened, leaflike, and curled away from the esophagus. The second or posterior salivary glands have exceptionally long and stout ducts. The glands themselves are long, stout anteriorly and taper posteriorly. They are very large. The esophagus is long and in its posterior third enters the crop which is composed almost in its entirety of an exceptionally large caecum free for about 4/s of its length. The short posterior esophagus leads into the typical bipartite stomach. The spiral caecum is large and strongly coiled. The digestive ducts lie sin- gularly close together and lead into the strongly bilobed digestive gland near the midline. The digestive gland is long and tubular. There are no surface indications of the digestive duct appendages which may, however, be bur- ied. The intestine is exceptionally long and tripartite. The anus bears two slender anal flaps. The ink sac is long, runs the length of the digestive gland 364 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Table 1. Measurements (in mm) and counts of 5 males of Octopus rapanui. Paratypes Holotype Mantle length 70 88 97 113 115 Mantle width 32 65 45 47 47 Head width 27 43 32 43 33 Arm length L R L R Ib, R L R I; R I 262-235 379 Ee 296, SOF S60 156 > 275 854 II 246" 3206 370) 359 290' 368 993537 373.) (4iSeG Ill 218 208 «©6328 7S 322 231 278) 240) Fee IV 170 186" 3325-3305 281 . 276.2219 7) 313209 Be 374 Total length 336 393 475 500 550 Arm width 11 jaja 12 14 14 Hect. Arm length 208 1D5r 231 240 279 Ligula length 3 1.8 go 7 9 Calmus length 1 0.45 3 2 3 Sucker diameter 7 9 11 12 10 Web depth 43 48.5 40 71 83 30 50 61 58 58 78 84 61 Til 73 51 49 82 48 80 58 70 58 80 66 31 4] 76 53) 38 62 50 60 78 3 Gill lamellae 11 12 12 ; 12 12 and is deeply set into it. The free duct is very long, strongly convoluted at the proximal end, then straight, but looped several times at the distal end. It enters the intestine just proximal to the anus. The female genitalia consist of a large round posterior ovary with paired oviducts and glands. The proximal oviducts are short and narrow. The oviducal glands are spherical with a dark band around the middle. The distal oviducts are long and a little stouter than the proximal ones. No mature eggs were seen. The male genitalia were dissected out of two specimens. The penis is stout with a long penial diverticulum and a short stout supplementary di- verticulum at the midpoint at the juncture with the duct from Needham’s sac (PLI 20-29). Needham’s sac is rather small but with a long accessory gland. Two of the males had spermatophores. These were long (SpLI 66-75), slender, with large sperm reservoirs (SpRI 42.3—43.4). There were no un- usual morphological features. The specimens were all in a poor state of preservation and little can be told concerning their original coloration. They appear to be somewhat flesh- colored ventrally with a darker purplish cast dorsally. The dorsal and lateral surfaces of the mantle, head and arms are covered with scattered but prom- VOLUME 92, NUMBER 2 365 Table 2. Measurements (in mm) and counts of 4 females of Octopus rapanui. Paratypes Mantle length 27 69 90 107 Mantle width 17 35 54 57 Head width 14 28 43 4] Arm length 1, R JL, R L R IG, R I 115 86 310 156 394 364 387 383 II 104 111 291 296 389 Sri 321 370 Ill 99 86 264 DT 380 77/5 193 381 IV 93 97 226 234 336 349 335 33) Total length 144 387 500 508 Arm width 6 9 14 16 Sucker diameter 4 7 10 10 Web depth sector A 17 49 69 68 B 23 21 57 47 68 69 38 29 E 19 US 53 34 55 i, 67 64 D 16 2D, 51 34 81 40 60 58 E 19 34 33 68 Gill lamellae 11 11 11 12 inent tubercles. Some of the tubercles over the eyes are somwhat larger than the others but not sufficiently so as to be called cirri. Type Locality. —Hanga Piko, Easter Island. Etymology.—This octopus is named for Easter Island which in the native language is called Rapa Nui. The specific name is, therefore, a noun in apposition. Discussion.—Octopus rapanui is unusual in a number of aspects and shows no close relationship with any of the other species of the genus in the Indo-Pacific. While it might be grouped with those species having long first arms such as Octopus macropus and O. ornatus, it differs in so many other features that this relationship cannot be seriously considered. One of the distinctive features is the secondary diverticulum of the penis. This is unusual in the octopods and is not known in other species of Octopus sensu stricto. Most of the other distinguishing characters are associated with the digestive tract. The straight spine-like rostrum of the lower beak is known so far only in Pareledone polymorpha (Robson, 1930) in which this feature is even more accentuated. Associated with this is the very long rachidian teeth of the radula which are longer (in the young specimen) than in any other known octopod. These two features suggest a specialized diet, perhaps barnacles or some type of bivalve. There are no indications of food in any of the stomachs or crops investigated. Other specialized features are the very long, slender but large second salivary glands, the comparatively enor- mous crop, elongate digestive gland, and the long tripartite intestine. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 366 —— i. a Se ee a eee eR eS cl Cl IT se[owel TID Cl cl AddOov Aqovad Vdd) Vdddo AVd4d) B[NULIOF GoM TI ATT Il AVI II ATI ATIVIVI AVIVIVI e[NUoy Way veP a = ; = = XOPUI IOAIOS31 “Wiads 97 SG = = == XOpUI YIpIM ‘ulIeds 99 SL =, = — XOpUl YJsUZ] ‘Wieds LG (G6 67 a 02 XOpUl Y}SUI] sIUDg Se 6°87 CAee = eee XOpuUl YJsUZ] snweyes One Gr 0'V = vl XOPUI YIZUZ] BNI] Iv $8 CL a 66 XOpUI We -1D90H] 6 IT IT Ol Ol [eWIOU XopuUl Jayons 0¢ €¢ CC (LG 61 xopul yidap qa cl cl al SC 91 XOpUI UIPIM Wiy 9L CL LL 96 6L XOpul Yjsue] WIy 8¢ el 9¢ eG LC XOpUl Wie suey 67 8€ (ele 6P 6£ XOpUl YIPIM peo Iv CY 5) 4 vL OV XOpul YIpPIM opuey] SII Ell L6 88 OL yi sug] spuey adAjo[oH I. 7 eS Ee ee ee ee ‘Inuvdps sndojIQ JO sojew ¢ Jo syuNod pue se[NuoJ “suons1odoid ATIpog Jo sooipuy = “¢ aquy, VOLUME 92, NUMBER 2 367 Table 4. Indices of bodily proportions, formulas and counts of 4 females of Octopus ra- panui. Mantle length 27 69 90 107 Mantle width index 63 Si 60 a5 Head width index 52 41 48 38 Mantle arm index 23 22 23 28 Arm length index 80 80 78 76 Arm width index 22. 13 15 15 Web depth index 22 18 21 18 Sucker index normal 15 10 11 9 Arm formula L..U1.1V L.I.I.1V L.IU.U1.1V L.I.I.1V Web formula BCDEA BCDAE DA;— BCE A = BCDE Gill lamellae 11 11 1. 12 Easter Island is a high island with steep-to shores where competition for food is probably intense; hence an adaptation to a favorable food niche would be more valuable and perhaps reinforced than in the more widespread coral reef habitats of the other areas of the tropical Pacific. While this species appears to be distinct, the description of any new species of Octopus from the Pacific must be approached with caution. Ex- ternal characters would not have sufficed to distinguish this species from others, but it should also be noted that we know little of the internal anatomy of Pacific octopods in general, and it is possible that future work may show this combination of characters in another named species. Until then it seems best to regard O. rapanui as a new and distinctive member of the Indo- Pacific octopod fauna. Acknowledgments I with to thank Dr. Harald A. Rehder of the Smithsonian Institution and Drs. Ian E. Efford and Jack Mathias of the University of British Columbia for the opportunity to study the material described above. The illustration of the whole animal is by Charles Messing; the remainder by Banjong Mian- manus and the writer. This work was supported by National Science Foundation grant DEB 76- 03456 which is gratefully acknowledged. This paper constitutes a scientific contribution from the Rosenstiel School of Marine and Atmospheric Science of the University of Miami. Literature Cited Robson, G. C. 1932. A monograph of the recent Cephalopoda based on the Collections in the British Museum (Natural History), Part II The Octopoda (excluding the Octopodinae).— The British Museum, London, pp. 1-359, 79 figs., 6 pls. Rosenstiel School of Marine and Atmospheric Science, University of Miami, 4600 Rickenbacker Causeway, Miami, Florida 33149. PROC. BIOL. SOC. WASH. 92(2), 1979, pp. 368-379 REVISION OF AMERICAN SPECIES OF THE MARINE AMPHIPOD GENUS PARAPHOXUS (GAMMARIDEA: PHOXOCEPHALIDAE) J. Laurens Barnard Abstract.—Most American species formerly placed in Paraphoxus Sars by J. L. Barnard (1960) are distributed into Metharpinia Schellenberg and the following 5 new genera: Eobrolgus, Eyakia, Foxiphalus, Grandifoxus and Rhepoxynius. New diagnoses parallel to those published for previously known phoxocephalid genera by Barnard and Drummond (1978) are given for all 6 American genera, plus Microphoxus. Metharpinia Schellenberg, revised Metharpinia Schellenberg, 1931:65.—J. L. Barnard, 1960:182. Diagnosis.—Eyes present. Flagella of antennae 1-2 unreduced in female; article 2 of antenna | ordinary to elongate, ventral setae narrowly to widely spread; article 1 of antenna 2 not ensiform, article 3 with 2 setules, facial spines on article 4 in 2 or more rows, article 5 ordinary in size. Right man- dibular incisor with 3 teeth; molar not triturative, medium, pillow-shaped or elongate, conical, weakly granulated, bearing 4 or more splayed, semi- articulate spines, not (type) or bearing pubescence; palpar hump small. Palp of maxilla 1 biarticulate; inner plate with 4 setae. Setation of maxilla 2 ordinary. Inner plate of maxillipeds ordinary; apex of palp weakly protu- berant, dactyl elongate, apical nail mostly immersed, obsolescent. Gnathopods ordinary, small, similar; article 5 of gnathopods 1-2 elongate, without eusirid attachment; palms oblique, hands ordinary, ovatorectan- gular, poorly setose anteriorly. Article 2 of pereopod 5 broad; articles 4—5 of pereopods 5-6 broad to medium; article 2 of pereopods 5—6 not setose posteriorly; pereopod 7 ordinary, article 2 naked or weakly setulose ven- trally, article 3 ordinary, dactyl normal. Peduncle of uropod 1 normally elongate, without apicoventral spike, with- out displaced apicomedial spine; peduncular apices of uropods 1-2 not combed; inner ramus of uropod 1 with one row of marginal spines, rami with one or two accessory nails but no main nails, inner ramus of uropod 2 ordinary; uropod 3 ordinary, article 2 of outer ramus long, carrying 2 medium apical setae. Telson extraordinary, with 1-3 apical spines or setae on each lobe plus setules, with special dorsal and lateral brush of setae. Epimera 1-3 bearing numerous long posterior setae, without midfacial setae above ventral facial ridge, epimeron 3 ordinary. Urosomite 1 with lateral facial setae, often bearing one or more midventral crescents or bundles of setae; urosomite 3 without dorsal hook or with weak dorsal hump. VOLUME 92, NUMBER 2 369 Description.—Rostrum fully developed, constricted; pubescence on ar- ticle 1 of antenna | in male present; calceoli on male primary flagellum of antenna | present, calceoli on article 5 of male antenna 2 present, flagellum in male with calceoli; prebuccal parts ordinary, poorly separated from each other, upper lip dominant, epistome without spike, occasionally with tiny cusp; right lacinia mobilis bifid, thin, article 1 of mandibular palp short, palp medium to thin, apex of article 3 oblique, article 2 without outer setae; lower lip bearing cones; outer plate of maxilla 1 with 11 spines, one spine especially thickened; inner plates of maxilliped especially thick, ordinarily setose; coxae 2—4 without special anterodorsal humps; all posterior spines on article 6 of pereopods 3—4 thick and stiff, midapical spine or seta present, article 2 of pereopod 7 without facial setae, article 6 especially densely spinose and digitate apically; peduncle of uropod | with dorsolateral spines confined apically, medial spines widely spread, peduncle of uropod 2 with one medial spine or setule confined apically or with spines widely spread; peduncle of uropod 3 lacking extra subapical setae or spines; telson with ordinary pair of midlateral or dorsal setules on each side. Type-species.—Metharpinia longirostris Schellenberg, 1931 (selected by Barnard and Drummond, 1978). Other species.—floridana (Shoemaker, 1933); jonesi (J. L. Barnard, 1963). . Relationship.—Easily distinguished from all other new American genera in the retention of subapical, supernumerary spination on one or more rami of uropods | and 2. This seemingly small attribute has been found to be of generic value by Barnard and Drummond (1978). Microphoxus J. L. Barnard Microphoxus J. L. Barnard, 1960:291. Diagnosis.—Eyes present. Flagella of antennae 1-2 unreduced in female; article 2 of antenna | ordinary, ventral setae widely spread; article 1 of antenna 2 not ensiform, article 3 with 2 setules, facial spines on article 4 in 2 or More rows, without special apical spines, article 5 ordinary in size. Right mandibular incisor with 3 teeth; molar not triturative, small, pillow- shaped, bearing 3—4 splayed, semiarticulate spines, not bearing pubescence; palpar hump small. Palp of maxilla 1 biarticulate, inner plate with 1-4 setae. Setation of maxilla 2 ordinary. Inner plate of maxillipeds ordinary, apex of palp not or weakly protuberant, dactyl elongate, apical nail distinct, short. Gnathopods small, similar, article 5 elongate, without eusirid attachment, palms almost transverse, hands heavily setose anteriorly, almost tricho- phoxin in shape. Article 3 of pereopod 5 of broad form, articles 4—5 of pereopods 5-6 broad, article 2 of pereopods 5-6 not setose posteriorly, pereopod 7 ordinary, article 2 naked or weakly setulose ventrally, article 3 ordinary, dactyl normal. 370 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Peduncle of uropod 1 normally elongate, without apicoventral spike, with- out special displaced spine, peduncular apices of uropods 1-2 not combed; inner ramus of uropod 1 with one row of marginal spines, one or more rami continuously spinose to apex, or with subapical spines (not of nail category) inner ramus of uropod 2 ordinary; uropod 3 ordinary, article 2 of outer ramus carrying 2 long apical setae. Telson ordinary, with only 1-2 apical spines, or setae on each lobe plus setules, with special dorsal and lateral spines or setae. Epimera 1-2 lacking or bearing numerous long posterior setae, without midfacial setae above ventral facial ridge, epimeron 3 ordi- nary, of rounded classification and bearing 3 or more long setae. Urosomite 1 with or without lateral facial setae, bearing one or more midventral cres- cents or bundles of setae, or generally naked except for sparse apicoventral setae or spines near base of uropod 1; urosomite 3 with dorsal hook or special process. Description.—Rostrum poorly developed, constricted; [? pubescence on article 1 of antenna 1 in male unknown, calceoli on male primary flagellum of antenna | unknown, calceoli on article 5 of male antenna 2 unknown, flagellum in male ?without calceoli]; prebuccal parts ordinary, poorly sep- arated from each other, both epistome and upper lip dominant; right lacinia mobilis bifid, thin, article 1 of mandibular palp short, palp medium, apex of article 3 oblique, article 2 without outer seate; lower lip lacking cones; outer plate of maxilla 1 with 11 spines, one spine especially thickened; inner plates of maxilliped ordinarily setose; coxae 2—4 without special anterodorsal humps; all posterior spines on article 6 of pereopods 3-4, thick and stiff, midapical spine or seta present; article 2 of pereopod 7 without facial setae; peduncle of uropod 1 with dorsolateral spines confined apically, medial spines confined apically, peduncle of uropod 2 with only one medial spine or setule confined apically; peduncle of uropod 3 lacking extra subapical setae or spines; telson with ordinary pair of midlateral or dorsal setules on each side. Type-species.—Microphoxus minimus J. L. Barnard, 1960 (original des- ignation). Other species.—Metharpinia cornuta Schellenberg, 1931. Relationship.—Microphoxus differs from Metharpinia in the pair of large dorsal teeth on urosomite 3, and in the retention of a dactylar nail on the maxilliped. These do not seem to be strong characters, but in Australian phoxocephalids they are used to separate genera (Barnard and Drummond, 1978). However, American phoxocephalids are not as strongly divisible into groups as are Australian phoxocephalids and with the low level of explo- ration in South America one may expect to discover species intermediate between Metharpinia and Microphoxus. The type-species is very convergent to the Brolginae although it clearly is descendent from the Birubiinae cluster of genera. The characters bringing VOLUME 92, NUMBER 2 371 it close to Brolginae are the presence of only 3 molarial spines, the presence of only one spine in the final row of facial spines on article 4 of antenna 2 and the reduction of setae on the inner plate of maxilla 1 (reduced to one). Obviously the Brolginae must be redefined or abolished, and this problem is now under study. Rhepoxynius, new genus Etymology.—From “‘rhepo,’’ slope and ‘‘oxyno,’’ sharpen. Masculine. Diagnosis.—Eyes present. Flagella of antennae 1—2 unreduced in female, though somewhat short on antenna 1; article 2 of antenna 1 ordinary to elongate, ventral setae widely to narrowly spread; article 1 of antenna 2 weakly ensiform, article 3 with 2 setules, facial spines on article 4 in 2 or more rows, article 5 ordinary in size. Right mandibular incisor with 3 teeth, molar not triturative, small, pillow-shaped, bearing 4 or more splayed, semi- articulate spines, usually bearing pubescence; palpar hump small. Palp of maxilla 1 biarticulate, inner plate with 4 setae. Setation of maxilla 2 ordi- nary. Inner plate of maxillipeds ordinary; apex of palp weakly protuberant, dactyl elongate, apical nail distinct to weak. Gnathopods ordinary, small, similar, article 5 of gnathopods 1-2 elongate, without eusirid attachment; palms weakly oblique to transverse, hands or- dinary, ovatorectangular to weakly trichophoxin in shape, poorly setose anteriorly. Article 2 of pereopod 5 of broad form; articles 4—5 of pereopods 5—6 broad to medium; article 2 of pereopods 5-6 not setose posteriorly; pereopod 7 ordinary, article 2 naked or weakly setulose ventrally, article 3 ordinary, articles 5-6 usually with weak apical comb, dactyl normal. Peduncle of uropod | normally elongate, without apicoventral spike, with or without displaced apicomedial spine, peduncular apices of uropods 1-2 combed or not, inner ramus of uropod 1 with one row of marginal spines, no rami continuously spinose to apex; inner ramus of uropod 2 ordinary; uropod 3 ordinary, article 2 of outer ramus carrying 2 long apical setae. Telson ordinary, with 2—4 apical spines or setae on each lobe plus setules, without special dorsal and lateral spines or setae. Epimera 1-2 lacking long posterior setae, without midfacial setae above ventral facial ridge, epimeron 3 ordinary, all posterior setae confined to narrow ventral clump. Urosomite 1 with or without facial setae or spines, bearing 2 ventral crescents or bun- dles of setae, otherwise generally naked except for sparse apicoventral setae or spines near base of uropod 1; urosomite 3 without dorsal hook or special process. Description.—Rostrum fully developed, constricted; medial fuzz on ar- ticle 1 of antenna | in male present; calceoli on male primarily flagellum of antenna 1 present, calceoli on article 5 of male antenna 2 absent, or un- known, flagellum in male with calceoli; prebuccal parts ordinary, poorly B72 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON separated from each other, epistome with or without spike, right lacinia mobilis bifid or simple, thin, article | of mandibular palp short, palp medium to thin, apex of article 3 oblique, article 2 with or without outer setae; lower lip bearing or lacking weak cones; outer plate of maxilla 1 with 9-11 spines, one spine especially thickened; inner plates of maxilliped ordinarily setose; coxae 2—4 without special anterodorsal humps; all posterior spines on article 6 of pereopods 3-4 thick and stiff, midapical spine or seta absent or vestigial, article 2 of pereopod 7 without facial setae; peduncle of uropod 1 with dorsolateral spines confined apically or widely spread, medial spines con- fined apically or widely spread, peduncle of uropod 2 with only one medial spine or setule confined apically, peduncle of uropod 3 lacking extra sub- apical setae or spines; telson with ordinary pair of midlateral or dorsal se- tules on each side. Type-species.—Pontharpinia epistoma Shoemaker, 1938. Other species.—abronius (J. L. Barnard, 1960); bicuspidatus (J. L. Bar- nard, 1960); daboius (J. L. Barnard, 1960); fatigans (J. L. Barnard, 1960); gemmatus (J. L. Barnard, 1969); heterocuspidatus (J. L. Barnard, 1960); lucubrans (J. L. Barnard, 1960); stenodes (J. L. Barnard, 1960); tridentatus (J. L. Barnard, 1954); variatus (J. L. Barnard, 1960); vigitegus (J. L. Bar- nard, 1971). Relationship.—This diverse American genus is the counterpart of Biru- bius Barnard and Drummond (1976) from Australia, but differs from it only in the ensiform process of antenna 2. At first, this cluster of species was believed to be divisible into 2 genera, but the second genus, at first characterized by the attributes of P. stenodes, is intergraded by several species. Rhepoxynius stenodes differs from R. _ epistomus in the bifid right lacinia mobilis with the appearance of two fused raker spines, the great reduction or loss in epistomal spike, the arrangement of facial setae on article 4 of pereopods 3—4 almost parallel to the apical margin (not spread axially), the loss of the medial displaced spine on the peduncle of uropod 1, the absence of molarial plumes, generally in the shorter flagella of antenna 1 with smaller aesthetascs and stouter setules, and in the wider spread of ventral setae on article 2 of antenna 1. But all of these characters find intermediates in several of the other species listed for this genus. The genus therefore has a wide diversity in uropod 1 armament whereas Birubius has a wide diversity in head shape. Foxiphalus, new genus Etymology.—Contrived phonetic name, masculine. Diagnosis.—Eyes present. Flagella of antennae 1-2 slightly reduced in female; article 2 of antenna 1 elongate, ventral setae widely to narrowly spread; article 1 of antenna 2 ensiform, article 3 with 2 setules, facial spines VOLUME 92, NUMBER 2 373 on article 4 in 2 or more rows, article 5 ordinary in size. Right mandibular incisor with 3 teeth, molar not triturative, granulate-striate, small to medi- um, or elongate plaque-forming, bearing 4 or more splayed, semiarticulate spines, usually bearing pubescence; palpar hump medium. Palp of maxilla 1 biarticulate, inner plate with 4 setae. Setation of maxilla 2 ordinary. Inner plate of maxilliped ordinary, apex of palp weakly protuberant, dactyl elon- gate, apical nail distinct, short. Gnathopods ordinary, small, similar, article 5 elongate, without eusirid attachment, palms oblique, hands of gnathopods 1-2 ovatorectangular, elon- gate, poorly setose anteriorly. Article 2 of pereopod 5 of broad form; articles 4-5 of pereopods 5—6 narrow to medium; article 2 of pereopods 5-6 not setose posteriorly; pereopod 7 ordinary, article 2 naked or weakly setulose ventrally, article 3 ordinary, dactyl normal. Peduncle of uropod 1 normally elongate, without apicoventral spike, with or without displaced enlarged apicomedial spine, peduncular apices of uro- pods 1-2 not combed, inner ramus of uropod | with one row of marginal spines, no rami continuously spinose to apex, inner ramus of uropod 2 ordinary; uropod 3 elongate, article 2 of outer ramus carrying 2 short apical setae. Telson ordinary, with only 2—4 apical spines or setae on each lobe plus setules, often with special dorsal and lateral spines or setae. Epimera (1), 2, 3, bearing numerous long posterior setae, without midfacial setae above ventral facial ridge, epimeron 3 ordinary. Urosomite 1 without large lateral facial spines, bearing one or more lateral or midventral crescents or bundles of setae; urosomite 3 without dorsal hook or special process. Description.—Rostrum fully developed, unconstricted; pubescence on article 1 of antenna | and articles 3—4 of antenna 2 in male present; calceoli on male primary flagellum of antenna 1 present, calceoli on article 5 of male antenna 2 present, flagellum in male with calceoli; prebuccal parts ordinary, poorly separated from each other, upper lip dominant; right lacinia mobilis bifid or simple, thin; article 1 of mandibular palp short, palp medium to thick, apex of article 3 oblique, article 2 occasionally with outer setae; lower lip bearing cones; outer plate of maxilla 1 with 11 spines, one spine espe- cially thickened; inner plates of maxilliped thick, ordinarily setose; coxae 2—4 without special anterodorsal humps; all posterior spines on article 6 of pereopods 3-4 thick and stiff, midapical spine or seta present, elongate; peduncle of uropod 1 with dorsolateral spines confined apically, medial spines widely spread, peduncle of uropod 2 with only one medial setule confined apically, peduncle of uropod 3 lacking extra subapical setae or spines; telson with ordinary pair of dorsal setules on each side. Type-species.—Pontharpinia obtusidens Alderman, 1936. Other species.—cognatus (J. L. Barnard, 1960); major (J. L. Barnard, 1960); similis (J. L. Barnard, 1960). Relationship.—This genus differs from Birubius in the presence of pos- 374 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON terior setae on epimeron 2 and the loss of the nail on the maxillipedal dactyl. It differs from Rhepoxynius in the unconstricted rostrum. Grandifoxus, new genus Etymology.—Contrived from root of type-species and contraction of phoxus. Masculine. Diagnosis.—Eyes present. Flagella of antennae 1—2 unreduced in female, article 2 of antenna 1 ordinary to elongate, ventral setae narrowly to widely spread; article 1 of antenna 2 not to strongly ensiform, article 3 with 3+ setules or setae in adults, facial spines on article 4 in 2 or more rows, lacking Special apical spines, article 5 ordinary in size. Right mandibular incisor with 3 teeth, molar not triturative, pillow-shaped, bearing 4 or more splayed, semiarticulate spines, usually bearing pubescence; palpar hump small to medium. Palp of maxilla 1 biarticulate, inner plate with 4 setae. Setation of maxilla 2 ordinary. Inner plate of maxillipeds ordinary, apex of palp not or weakly protuberant, dactyl elongate, apical nail mostly immersed, short. Gnathopods ordinary, small, similar, gnathopod 2 weakly enlarged, article 5 of gnathopods 1-2 elongate, without eusirid attachment, palms oblique to transverse, hands of gnathopods 1-2 setose anteriorly, weakly trichophoxin in shape. Article 2 of pereopod 5 of broad form; articles 4—5 of pereopods 5-6 broad; article 2 of pereopods 5—6 not setose posteriorly; pereopod 7 ordinary, article 2 naked ventrally, article 3 ordinary, dactyl normal. Peduncle of uropod | normally elongate, without apicoventral spike, with or without enlarged displaced medial spine, peduncular apices of uropods 1-2 not combed, inner ramus of uropod 1 with one row of marginal spines, no rami continuously spinose to apex, inner ramus of uropod 2 ordinary; uropod 3 ordinary, very short, article 2 of outer ramus carrying 2 medium to long apical setae. Telson ordinary, with 2—4 apical spines or setae on each lobe plus setules, usually with special dorsal and lateral spines or setae. Epimera (1), 2, 3 bearing numerous long posterior setae, without midfacial setae above ventral facial ridge, epimeron 3 ordinary. Urosomite | without large lateral facial spines, bearing one or more midventral or lateral cres- cents or bundles of setae; urosomite 3 without dorsal hook or special pro- cess. Description.—Rostrum fully developed, constricted; pubescence on ar- ticle 1 of antenna 1 in male present; calceoli on male primary flagellum of antenna | present, calceoli on article 5 of male antenna 2 present, flagellum in male with calceoli; prebuccal parts ordinary, poorly separated from each other, both epistome and upper lip dominant; right lacinia mobilis bifid or simple, flabellate or thin, article 1 of mandibular palp short, palp medium, apex of article 3 oblique, article 2 with or without outer setae; lower lip bearing cones; outer plate of maxilla 1 with 11 spines, one spine especially VOLUME 92, NUMBER 2 375 thickened; inner plates of maxilliped ordinarily setose; coxae 2—4 with spe- cial apical humps; all posterior spines on article 6 of pereopods 3—4 thick and stiff, midapical spine or seta present; peduncle of uropod 1 with dor- solateral spines confined apically, medial spines widely spread, peduncle of uropod 2 with only one medial spine or setule confined apically, or rarely with widely spread spines; peduncle of uropod 3 lacking extra subapical setae or spines; telson with ordinary pair of midlateral or dorsal setules on each side. Type-species.—Phoxus grandis Stimpson, 1856 (=Pontharpinia milleri Thorsteinson, 1941). Other species.—longirostris (Gurjanova, 1938, 1951); robustus (Gurja- nova, 1938, 1951). Relationship.—Differing from Birubius, Rhepoxynius and Foxiphalus in the presence of more than 2 lateral setae on article 3 of antenna 2 and the humped coxae. Also characterized by very short article 2 on outer ramus of uropod 3 and, for the most part, supernumerary telsonic spines. Eyakia, new genus Etymology.—Named for a group of North American Indians. Feminine. Diagnosis.—Eyes present. Flagella of antennae 1-2 unreduced in female; article 2 of antenna 1 ordinary, ventral setae widely spread; article 1 of antenna 2 not ensiform, article 3 with 2 setules, facial spines on article 4 in 2 rows, plus special apical spines, article 5 ordinary in size. Right mandib- ular incisor with 3 teeth, molar not triturative, pillow-shaped, bearing 3—4 splayed, semiarticulate spines, one of these very large, usually bearing pu- bescence; palpar hump small. Palp of maxilla 1 biarticulate, inner plate with 4 setae. Setation of maxilla 2 ordinary. Inner plate of maxillipeds ordinary, apex of palp not or weakly protuberant, dactyl elongate, apical nail distinct, short. Gnathopods dissimilar, gnathopod 2 moderately to strongly enlarged, ar- ticle 5 of gnathopod 1 of ordinary length, but short on gnathopod 2, without eusirid attachment, palms oblique; hands of gnathopods 1-2 narrowly ovate, elongate, poorly setose anteriorly; article 2 of pereopod 5 of broad form, but tapering distally; articles 4-5 of pereopods 5—6 narrow to medium; ar- ticle 2 of pereopods 5-6 not setose posteriorly; pereopod 7 oridnary, article 2 naked or weakly setulose ventrally, article 3 ordinary, dactyl normal. Peduncle of uropod 1 normally elongate, without apicoventral spike, with- out enlarged displaced spine, peduncular apices of uropods 1—2 not combed; inner ramus of uropod | with one row of marginal spines, no rami contin- uously spinose to apex; inner ramus of uropod 2 ordinary, uropod 3 ordi- nary, article 2 of outer ramus carrying 2 medium to long apical setae. Telson ordinary, with only 1-2 apical spines or setae on each lobe plus setules, 376 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON without special dorsal and lateral spines or setae. Epimera 1—2 lacking long posterior setae, without midfacial setae above ventral facial ridge, epimeron 3 ordinary. Urosomite 1 without large lateral facial spines, bearing one or more ventral crescents or bundles of setae or generally naked except for Sparse apicoventral setae or spines near base of uropod 1, urosomite 3 without dorsal hook or special process. Description.—Rostrum fully developed, unconstricted, pubescence on article 1 of antenna | in male present; calceoli on male primary flagellum of antenna | present, calceoli on article 5 of male antenna 2 present, flagellum in male with calceoli; prebuccal parts ordinary, poorly separated from each other, both epistome and upper lip dominant; right lacinia mobilis bifid, flabellate; article 1 of mandibular palp short, palp medium, apex of article 3 oblique, article 2 without outer setae; lower lip bearing cones; outer plate of maxilla 1 with 11 spines, one spine especially thickened; inner plates of maxilliped ordinarily setose; coxae 2—4 without special anterodorsal humps; all posterior spines on article 6 of pereopods 1-2 thick and stiff, midapical Spine or seta present; article 2 of pereopod 7 without facial setae; peduncle of uropod | with dorsolateral spines confined apically, medial spines or setae widely spread, peduncle of uropod 2 with only one medial spine or setule confined apically, peduncle of uropod 3 lacking extra subapical setae or spines; telson with ordinary pair of midlateral or dorsal setules on each side. Type-species.—Parharpinia calcarata Gurjanova, 1938, 1951. Other species.—robustus (Holmes, 1908); subuncigerus (Kudrjaschov, 1965); uncigerus (Gurjanova, 1938, 1951). Relationship.—Similar to Parharpinia Stebbing and Protophoxus K. H. Barnard but differing from both in the absence of displaced peduncular spine on uropod 1 and in the enlarged third spine on the mandibular molars. The latter character distinguishes Eyakia also from Birubius Barnard and Drum- mond; in addition Eyakia differs from Birubius in the tapering article 2 of pereopod 5. Eobrolgus, new genus Etymology.—From ‘‘eos’’ early and Brolgus, an Australian genus; mas- culine; noting Eobrolgus more plesiomorphic than Brolgus in uropod 3. Diagnosis.—Eyes present. Flagella of antennae 1—2 unreduced in female; article 2 of antenna 1 ordinary, ventral setae confined apically; article 1 of antenna 2 not ensiform, article 3 with 2 setules, facial spines on article 4 in 2 or more rows, lacking special apical spines, article 5 ordinary in size. Right mandibular incisor with 3 teeth, molar not triturative, pillow-shaped, bearing 4 or more splayed, semiarticulate spines, usually bearing pubes- cence, palpar hump medium. Palp of maxilla 1 biarticulate, inner plate with VOLUME 92, NUMBER 2 S77 4 setae. Setation of maxilla 2 ordinary. Inner plate of maxillipeds ordinary, apex of palp not or weakly protuberant, dactyl elongate, apical nail obso- lescent to absent. Gnathopods small, similar, article 5 of ordinary length, very short, cryptic on gnathopod 2, elongate on gnathopods 1, 2, palms oblique, hands of gnathopods 1-2 ovatorectangular, poorly setose anteriorly. Article 2 of per- eopod 5 of broad form; articles 4-5 of pereopods 5-6 broad and narrow respectively, article 2 of pereopods 5-6 not setose posteriorly; pereopod 7 ordinary, article 2 naked ventrally, article 3 ordinary, dactyl normal. Peduncle of uropod 1 normally elongate, without apicoventral spike, with- out enlarged displaced spine, peduncular apices of uropods 1—2 combed; inner ramus of uropod 1 with one row of marginal spines, no rami contin- uously spinose to apex; inner ramus of uropod 2 ordinary, uropod 3 ordinary, article 2 of outer ramus, carrying 2 medium to long apical setae. Telson ordinary, with 1-2 apical spines or setae on each lobe plus setules, without special dorsal and lateral spines or setae. Epimera 1-2 lacking numerous long posterior setae, without midfacial setae above ventral facial ridge, epimeron 3 ordinary, bearing one or more long setae. Urosomite 1 without large facial spines, generally naked except for sparse apicoventral setae or spines near base of uropod 1; urosomite 3 without dorsal hook or special process. Description.—Rostrum fully developed, unconstricted, pubescence on article 1 of antenna | in male present; calceoli on male primary flagellum of antenna | present, calceoli on article 5 of male antenna 2 absent, flagellum in male with calceoli; prebuccal parts poorly separated from each other, both epistome and upper lip dominant; right lacinia mobilis bifid or simple, thin, article 1 or mandibular palp short, palp medium, apex of article 3 oblique, article 2 without outer setae; lower lip bearing cones; outer plate of maxilla 1 with 9 spines, one spine especially thickened; inner plates of maxilliped ordinarily setose; coxae 2-4 without special anterodorsal humps; all posterior spines on article 6 of pereopods 1-2 thick and stiff, midapical spine or seta present; article 2 of pereopod 7 without facial setae; peduncle of uropod 1 with dorsolateral spines confined apically, medial spines widely spread, peduncle of uropod 2 with only one medial spine or setule confined apically, peduncle of uropod 3 bearing extra subapical setae or spines; telson with ordinary pair of midlateral or dorsal setules on each side. Type-species.—Paraphoxus spinosus Holmes, 1905. Other species.—?pontarpioides (Gurjanova, 1953). Relationship.—Because of the short article 2 of antenna 1 and the place- ment of the major setal group at the apex of the article (in contrast to being spread out along the ventral margin), this genus has close affinities with the Brolginae as described by Barnard and Drummond (1978). In that subfamily 378 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON it keys closest to Paraphoxus Sars but differs from that genus in the pres- ence of more than 3 spines on the molars, the presence of large, or long, or numerous setae on epimeron 3, the presence of more than 2 setae on the inner plate of maxilla 1, the presence of a main apical spine on the inner plate of the maxilliped, and the division of the spines on article 4 of antenna 2 peduncle into several distinct groups the most proximal of which has more than 1 spine (Eobrolgus usually has 5). Eobrolgus thus is a very close mimic of Paraphoxus and superficially appears to be a brolgin but is not because of the characters of mandible and antenna 2. Instead, it belongs either to the Birubiinae or Parharpiniinae and differs from Birubius in the apical placement of setae on article 2 of antenna 1 or from other parharpiniins in the absence of supernumerary telsonic spination and untapering article 2 of pereopod 5. Literature Cited Alderman, A. L. 1936. Some new and little known amphipods of California.—University of California Publications in Zoology 41:53-74, figs. 1-51. Barnard, J. L. 1954. Marine Amphipods of Oregon.—Oregon State Monographs, Studies in Zoology. 8:1—103, 1 figure, plates 1-33. . 1960. The amphipod family Phoxocephalidae in the eastern Pacific Ocean, with anal- yses of other species and notes for a revision of the family. lies Hancock Pacific Expeditions 18:175—368, pls. 1-75, 1 chart. 1963. Relationship of benthic Amphipoda to invertebrate communities of inshore sublittoral sands of southern California.—Pacific Naturalist 3:437—467, figs. 1-7. . 1969. A biological survey of Bahia de Los Angeles, Gulf of California, Mexico, IV. Benthic Amphipoda (Crustacea).—Transactions San Diego Society Natural History 15:175—228, figs. 1-30. . 1971. Gammaridean Amphipoda from a deep-sea transect off Oregon.—Smithsonian Contributions to Zoology 61:1-86, figs. 1-48. , and M. M. Drummond. 1976. Clarification of five genera of Phoxocephalidae (marine Amphipoda).—Proceedings of the Biological Society of Washington 88:515—547, figs. 1-4. , and . 1978. Gammaridean Amphipoda of Australia, part 3. The Phoxocephal- idae.—Smithsonian Contributions to Zoology 245:1—-551, pls. 1-269. Gurjanova, E. 1938. Amphipoda, Gammaroidea of Siaukhu Bay and Sudzuhke Bay (Japan Sea).—Reports of the Japan Sea Hydrobiological Expedition of the Zoological Institute of the Academy of Sciences, USSR in 1934, 1:241-404, figs. 1-59 (in Russian with English title and summary). . 1951. Bokoplavy morei SSSR i sopredel’nyx vod (Amphipoda-Gammaridea).—Opre- deliteli po Faune SSSR 41:1—1031, figs. 1-705. —. 1953. Novye dopolneniya k dal’nevostochnoi faune morskik bokoplavov.—Trudy Zoologicheskogo Instituta 13:216—241, figs. 1-19. Holmes, S. J. 1905. The Amphipoda of southern New England.—Bulletin of the U.S. Bureau of Fisheries 24:459-529, pls. 1-13, numerous text figs. [unnumbered]. 1908. The Amphipoda collected by the U.S. Bureau of Fisheries Steamer, **Alba- tross,’’ off the west coast of North America, 1903 and 1904, with descriptions of a new family and several new genera and species.—Proceedings of the United States National Museum 35:489—543, figs. 1-46. VOLUME 92, NUMBER 2 pe Kudrjaschov, V. A. 1965. Novye vidy bokoplavov (Amphipoda, Gammaridea) iz vostochnoi chasti Oxotskogo Morja.—Zoologicheskii Zhurnal 44: 1776-1789, figs. 1-10. Schellenberg, A. 1931. Gammariden und Caprelliden des Magellangebietes, Siidgeorgiens und der Westantarktis.—Further Zoological Results of the Swedish Antarctic Expedition 1901-1903, 2(6): 1-290, figs. 1-136, pl. 1. Shoemaker, C. R. 1933. Amphipoda from Florida and the West Indies.—American Museum Novitates 598:1—24, figs. 1-13. . 1938. Two new species of amphipod crustaceans from the east coast of the United States.—Journal of the Washington Academy of Sciences 28:326-—332, figs. 1-2. Stimpson, W. 1856. On some California Crustacea.—Proceedings of the California Academy of Science 1:87—90. Thorsteinson, E. D. 1941. New or noteworthy amphipods from the North Pacific coast.— University of Washington Publications in Oceanography 4:50-96, pls. 1-8. Department of Invertebrate Zoology, Smithsonian Institution, Washing- ton, D.C. 20560 (Mailing address = NHB 163-W-323). PROC. BIOL. SOC. WASH. 92(2), 1979, pp. 380-383 NANNOSQUILLA VASQUEZI, A NEW STOMATOPOD CRUSTACEAN FROM THE ATLANTIC COAST OF PANAMA Raymond B. Manning Abstract.—The eighth Atlantic species of Nannosquilla is described from material collected in shallow water habitats at Punta Galeta, Atlantic coast of Panama. Species of Nannosquilla differ from other stomatopods in meth- ods of egg brooding. In 1977, Rafael Vasquez M., then with the Station Marine d’Endoume, Marseilles, forwarded a small collection of stomatopods taken during an eco- logical survey of the Punta Galeta [09°24’N, 79°52'’W] area on the Atlantic coast of Panama. Most of the specimens belong to an undescribed species of Nannosquilla, named below for Mr. Vasquez. All of the specimens reported below have been deposited in the National Museum of Natural History, Smithsonian Institution, Washington (USNM). I thank Mr. Vasquez for forwarding these specimens for study and M. Reaka, University of Maryland, for her comments on egg deposition in Nannosquilla and for reading the manuscript. The illustrations were pre- pared by my wife Lilly. Nannosquilla vasquezi, new species Figure 1 Material.—Panama, Atlantic coast, near Punta Galeta laboratory, Smith- sonian Tropical Research Institute; 09°23.9'N, 79°51.81'W; in Halodule wrightii bed often exposed at low tide; R. Vasquez M., leg.: Sta. PCS Ic, 1 2 TL (total length) 23.5 mm; Sta. PCS 1d, 1 ¢ TL 20 mm; Sta. PCS lg, 1 2 TL 21 mm; with ova: Sta. PCS 2c, f 2 TE 21 mm: Sta; PCS Saggiec TL 18.5 mm; Sta. PCS 3b, 1 6 TL 17.5 mm, 1 2 TL 20 mm. The male from Sta. PCS 3a is the holotype (USNM 171349); the other specimens are para- types. Diagnosis.—Size small, total lengths of adults 17.5 to 23.5 mm. Cornea subglobular, set distally on stalk and slightly overhanging stalk laterally. Ocular scales erect, subquadrate, bases fused medially. Rostral plate quad- rangular, lateral margins convex, subparallel, anterolateral angles rounded, unarmed, apex an abtuse point. Dactylus of claw with 7-9, usually 8, teeth, outer margin with 2 low prominences proximally, distalmost sharper. Man- dibular palp absent. 4 epipods present. Sixth abdominal somite produced VOLUME 92, NUMBER 2 381 Fig. 1. Nannosquilla vasquezi, new species, female paratype, TL 23.5 mm; a, Anterior part of body; b, Sixth abdominal somite, telson and uropod; c, Telson, ventral view; d, Uropod, ventral view (setae omitted). posterolaterally into sharp points. Telson broader than long, false eave with obtuse median projection only, submedian margins slightly concave, later- almost marginal teeth partially or completely visible in dorsal view. Marginal armature consisting of (on either side of midline) 6—10 (usually 8—9) sub- median denticles, mesials higher than laterals, 1 movable submedian tooth, adjacent to outermost denticle, and 3—4 (usually 4) fixed lateral teeth. Uro- pod with 2-3 stiff setae distally on inner margin and 4—S (usually 5) movable spines, distal 3 spatulate, on outer margin of proximal segment of exopod. Inner spine of basal prolongation of uropod the longer. Color.—Stellate brown chromatophores scattered over body, arranged in a line on posterior margins of thoracic and abdominal somites as well as along lateral margins of abdominal pleura. 2—3 pairs of larger spots present along gastric grooves. Size.—Total lengths of males 17.5 to 20 mm, of females 20—23.5 mm. Ova 382 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON found loose in vial with female from Sta. PCS lg were large, measuring 0.8 x 1.0 mm. Remarks.—This new species is very similar to N. hancocki (Manning, 1961), known from two localities off Venezuela: Isla Cubagua [10°49’N, 64°11’W], the type-locality, in 4-9 m, and off Isla de Margarita [11°00’N, 64°00’ W], in 38-40 m (Manning, 1969:76). Nannosquilla vasquezi has less divergent lateral margins on the rostral plate, fewer teeth, 7-8 rather than 9-10, on the claw, and further differs in several aspects of telson structure: the false eave has only a median projection, the movable submedian teeth are adjacent to rather than anterior to the outer submedian denticles, and there are 3—4 rather than 5 fixed lateral teeth. Only two of the specimens of N. vasquezi examined had more than 3 fixed teeth lateral to the movable submedians and those specimens each had 3 teeth on one side, 4 on the other. Habitat preferences of the two species may also be different. Nanno- squilla vasquezi was collected in beds of Halodule wrightii near the low tide line. Mr. Vasquez noted that the part of the beds in which the specimens were found usually is exposed at low tide. Nannosquilla hancocki has been taken in depths of 4—9 m and 38—40 m, but not intertidally. Manning (1969:69-83) provided accounts of the species of Nannosquilla than known from the western Atlantic. In the key to species given there (p. 71) N. vasquezi would key out with N. hancocki. Two additional western Atlantic species have been described since then (Manning, 1970). In the vial with the female from Sta. PCS lg were several loose ova, assumed to be the ova of this species. They were not formed into a ball. Stomatopoda are generally believed to carry their ova in a loose ball until the eggs hatch (Manning and Provenzano, 1963), but this is the third time that loose ova have been associated with a species of Nannosquilla. The first report was by Rathbun (1910:566) in her original account of N. decem- spinosa from Peru, who noted that it was found ‘“‘Living in vertical holes in the muddy sand of the inside beach at Capon. Small yellow eggs were often noted attached to the sides of the holes.’ The second observation was reported by Manning (1967:149) in the original account of N. anomala from San Clemente Island, California, where it was noted that eggs were found in some of the burrows. That some of the lysiosquillids differ from the squillids and gonodactylids in methods of egg care as well as in developmental stages (Gurney, 1946) suggests that the lysiosquillids comprise a fundamentally different stock from the other stomatopods. This could have wide ranging implications in stomatopod classification; the status of the Lysiosquillidae should now be investigated in light of these findings. VOLUME 92, NUMBER 2 383 Literature Cited Gurney, R. 1946. Notes on stomatopod larvae.—Proc. Zool. Soc. London 116(1):133-175, figs. 1-14. Manning, Raymond B. 1967. Nannosquilla anomala, a new stomatopod crustacean from California.—Proc. Biol. Soc. Washington 80:147-150, figs. 1-4. . 1969. Stomatopod Crustacea of the western Atlantic.—Stud. trop. Oceanogr. Miami 8:vili + 380 pp. 1970. Nine new American stomatopod crustaceans.—Proc. Biol. Soc. Washington 83:99-114. Manning, Raymond B., and Anthony J. Provenzano, Jr. 1963. Early larval stages of Gono- dactylus oerstedii Hansen. Studies on development of stomatopod Crustacea, I.—Bull. Mar. Sci. Gulf & Carib. 13(3):467—487. Rathbun, M. J. 1910. The stalk-eyed Crustacea of Peru and the adjacent coast.—Proc. U.S. Natl. Mus. 38(1766):531-620. Department of Invertebrate Zoology, National Museum of Natural His- tory, Smithsonian Institution, Washington, DC 20560. PROC. BIOL. SOC. WASH. 92(2), 1979, pp. 384-388 REDESCRIPTION OF BRUUNILLA NATALENSIS HARTMAN (POLYCHAETA: POLYNOIDAEB), ORIGINALLY REFERRED TO FAUVELIOPSIDAE Marian H. Pettibone Abstract.—The abyssal polychaete Bruunilla natalensis Hartman from the Mozambique Basin, originally placed in the Fauveliopsidae, is rede- scribed and referred to the Polynoidae: Macellicephalinae. The abyssal polychaete Bruunilla natalensis was originally described by Hartman (1971:1411) from a single small specimen collected by the R/V Anton Bruun in the Mozambique Basin and referred to her new family Fauveliopsidae. In his Keys to the Polychaete Families and Genera, Fau- chald (1977:118) characterized the genus Bruunilla as ‘‘Incertae Sedis, with- out obvious familial affiliations.’’ The description and figures by Hartman suggested to me that it might belong to the Polynoidae: Macellicephalinae, a group of deep and cold water polynoid species that I studied recently (Pettibone, 1976). A study of the holotype, which was kindly sent on loan from the Allan Hancock Foundation by Dr. Kristian Fauchald, revealed that the species does indeed belong to the Polynoidae. It should be pointed out that the single specimen was defective in a number of respects, as is often the case in abyssal soft-bodied polychaetes: all the setae and elytra, as well as most of the dorsal cirri were missing; the muscular pharynx was not extended and the jaws were not observed. The description of the species is Supplemented as much as the defective holotype will allow. I thank my colleague Meredith Jones for his review of this manuscript. Family Polynoidae Malmgren Subfamily Macellicephalinae Hartmann-Schroder, emended Pettibone, 1976 Genus Bruunilla Hartman, 1971, emended Type-species.—Bruunilla natalensis Hartman, 1971, by monotypy. Gen- der: feminine. Diagnosis.—Body short, flattened, tapered; segments 18 (first achaetous). Elytra and small elytrophores, emerging near bases of notopodia (similar in position to cirrophores of dorsal cirri on cirrigerous segments), 8 pairs on segments 2, 4, 5, 7, 9, 11, 13 and 15, with dorsal cirri on posterior 3 seg- ments. Prostomium oval, slightly bilobed, without frontal filaments or lateral VOLUME 92, NUMBER 2 385 Aas 63 ie —_ \\ \ LOmm ats Fig. 1. Bruunilla natalensis, holotype (AHF 1202): a, Dorsal view of anterior end; left parapodium of segment 3 had been cut off; styles of median antenna, right dorsal tentacular cirrus, right buccal cirrus of segment 2, right dorsal cirrus of segment 3 and elytra of segment 2 missing; b, Ventral view of same; left margin of wing-like structure overlapping segment 3 damaged. 386 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 0.5mm Fig. 2. Bruunilla natalensis, holotype (AHF 1202): a, Left cirrigerous parapodium from segment 3, anterior view; acicula dotted; b, Right cirrigerous parapodium from segment 8, anterior view; style of dorsal cirrus missing; c, One of jaws. antennae; ceratophore of median antenna inserted on middle of prostomium; paired palps long, filiform; without eyes. First or tentacular segment fused to prostomium; achaetous tentaculophores with 2 pairs of tentacular cirri lateral to prostomium; facial tubercle on upper lip trilobed. Segment 2 with buccal cirri attached to basal parts of parapodia, lateral to ventral mouth and lower lip; styles longer than following ventral cirri. Unique wing-like structure on ventral side of lower lip and projecting posteriorly on segment 3. Parapodia biramous, both rami with elongate acicular lobes; notopodia VOLUME 92, NUMBER 2 387 shorter than neuropodia. Setae unknown (all missing). Dorsal cirri with short cylindrical cirrophores and moderately long styles, bulbous basally; ventral cirri short, attached near distal ends of neuropodia. Without distinct dorsal tubercles on cirrigerous segments. Nephridial papillae indistinct, none enlarged. Pygidium oval, truncate, with dorsal anus and pair of anal cirri (?). Pharynx with papillae (number?) and 2 pairs of jaws; jaw plates with inner row of teeth. Bruunilla natalensis Hartman Figures 1, 2 Bruunilla natalensis Hartman, 1971:1411, fig. 3a—f.—Fauchald, 1977:118. Material examined.—Mozambique Basin, off Natal, southeast Africa, 34°06-10’S, 41°14—15’E, 4886-5069 m, red-brown mud, R/V Anton Bruun Sta. 382C, 31 August 1964—holotype (AHF 1202). Description.—The holotype and only known specimen has a length of 9 mm, a width of 3.5 mm, including the parapodia, and 18 segments, the last one very small. The body is flattened, tapered posteriorly, with the seg- mental lines poorly indicated (Fig. 3a, in Hartman). The elytra are all miss- ing; their elytrophores are rather small but slightly larger than the cirro- phores of the dorsal cirri, at least on the more anterior segments. Dorsal tubercles on the cirrigerous segments are indistinct. The anterior end was not clearly described and figured by Hartman (Fig. la, b; Fig. 3a, b, in Hartman). The oval prostomium is slightly bilobed, without lateral anten- nae; the clavate ceratophore of the median antenna is inserted near the middle of the prostomium; the style is now missing (long, filiform, as ob- served by Hartman); the palps, emerging from the anterior-lateral borders of the prostomium, have distinct palpophores and long filiform styles. The achaetous tentaculophores of segment 1 are fused basally to the lateral sides of the prostomium; they bear 2 pairs of tentacular cirri which are subequal in length to the median antenna and palps. Thus, there are 7 anterior filiform head appendages, not 5, as shown by Hartman (Fig. 3a, b); the facial tu- bercle on the upper lip, anterior to the ventral mouth, has 3 rounded lobes; the lateral lips are bulbous. Segment 2 (first setigerous segment) forms the lower lip, with a pair of long buccal cirri inserted ventrally on the basal parts of the parapodia; a pair of thickened bulbous lobes on the lower lip, with a median groove between them, projects freely on segment 3 and forms a flattened wing-like structure, tapering laterally (Fig. 1b; Fig. 3b, in Hart- man and referred to as ‘‘a pair of palplike flat pads’’). The biramous parapodia are elongated, about as long as the width of the body (Figs. la, b, 2a, b; Fig. 3a—3b, in Hartman). Both rami have stout acicula, tapering distally to slender hooked tips within projecting acicular lobes; the notopodia are smaller and shorter than the neuropodia. The setae 388 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON are all missing but there is evidence that there was a small bundle of no- tosetae and a fan-shaped bundle of neurosetae. The dorsal cirri have short cylindrical cirrophores; the styles are inflated basally and may extend be- yond the distal tips of the neuropodia; they are mostly missing (present on segments 3, left 6 and right 12; Fig. 2a, b; Fig. 3a, d, e, in Hartman). The ventral cirri are attached rather far distally and are short and tapered. The elytra are all missing but, judging from the small size of the elytrophores, they are probably small. It is difficult to distinguish elytrophores and cir- rophores in the posterior region but there appear to be 8 pairs of elytra, on segments 2, 4, 5, 7, 9, 11, 13, 15, with dorsal cirri on the last 3 small segments. The posterior end is somewhat damaged; the truncate pygidium, with the dorsal anus, extends beyond the small posterior parapodia. Nephridial pa- pillae are indistinct. The muscular pharynx, occupying an inflated area in segments 4-7, was not extended. A slit on the ventral side of the body revealed that there are 2 pairs of amber-colored jaws and typical large ve- sicular papillae around the opening (exact number not determined); one of the jaws was removed and showed a row of about 12 teeth along the inner border (Fig. 2c). Remarks.—Among the genera and species of Macellicephalinae covered by Pettibone (1976), Bruunilla natalensis is closest to Bathyvitiazia Petti- bone, with type-species B. pallida (Levenstein, 1971) from the Northwest Pacific off Mexico, in 3816 m. The latter species lacks the unique wing-like structure on the ventral side of segments 2 (lower lip) and 3; the upper lip has a bilobed oral lobe, instead of a trilobed facial tubercle; the border of the jaw plate has about 32 small teeth, instead of about 12 larger teeth; the styles of the dorsal cirri are long and tapering, instead of shorter and bulbous basally. The setae cannot be compared, since they are unknown in Bruunilla natalensis. Literature Cited Fauchald, K. 1977. The polychaete worms: definitions and keys to the orders, families and genera.—Nat. Hist. Mus. Los Angeles Co. Sci. Ser. 28:1—190. Hartman, O. 1971. Abyssal polychaetous annelids from the Mozambique Basin off southeast Africa, with a compendium of abyssal polychaetous annelids from world-wide areas.— Jour. Fish. Res. Bd. Canada, 28:1407-—1428. Levenstein, R. J. 1971. [Polychaete worms of the genus Macellicephala and Macellicepha- loides (Family Aphroditidae) from the Pacific Ocean.—/n Fauna of the Kurile-Kam- chatka Trench.] Trudy Institut Okeanologii P. P. Shirshov Akademiia Nauk SSSR, 92:18— 35. [In Russian, English Summary. ] Pettibone, M. H. 1976. Revision of the genus Macellicephala McIntosh and the subfamily Macellicephalinae Hartmann-Schréder (Polychaeta: Polynoidae).—Smithsonian Con- trib. Zool. 229:1-71. Department of Invertebrate Zoology, Smithsonian Institution, Washing- ton, D.C. 20560. : PROC. BIOL. SOC. WASH. 92(2), 1979, pp. 389-393 FRESHWATER TRICLADS (TURBELLARIA) OF NORTH AMERICA. XI. PHAGOCATA HOLLERI, NEW SPECIES, FROM A CAVE IN NORTH CAROLINA Roman Kenk Abstract.—Phagocata holleri, n. sp., from Mt. Jefferson Cave in Mt. Jefferson State Park, North Carolina, is described. The species resembles P. angusta of West Virginia by its external appearance, but differs from it conspicuously in the structure of its copulatory apparatus. The Appalachian region of the southeastern United States contains sev- eral unpigmented (white) species of the genus Phagocata. Some of these species show prominent auricular projections (P. bursaperforata Darlington from Georgia and Alabama and at least two undescribed species from North Carolina and Tennessee), while others lack such projections or auricles and may easily be confused in the field with the more widely distributed P. morgani (Stevens and Boring). Dr. Cato O. Holler, Jr. of the North Carolina Cave Survey forwarded to me several live planarians collected in a cave in Mt. Jefferson State Park, North Carolina. They proved to be a new species. Phagocata holleri, new species Type-material.—Holotype, whole mount of anterior end and sagittal sec- tions of the rest of the body on 6 slides, deposited in the National Museum of Natural History, Smithsonian Institution (USNM 57139); paratypes, sag- ittal sections of 3 specimens on 16 slides (USNM 57140-57142). External features (Figs. 1A, 1B).—The body is very slender, measuring at maturity up to 15 mm in length and | mm in width. The anterior end of the quietly gliding animal is truncate, with a straight or slightly concave frontal margin and laterally projecting rounded edges. Behind the head, the body first narrows, then gradually widens again to maintain its width close to the posterior end. The body is unpigmented, white. The head bears a pair of very small eyes, barely noticeable under the dissecting microscope (they do not show in the photograph of the specimen in Fig. 1A). They are placed close together at a mutual distance of perhaps '!/5; the width of the body at the level of the eyes and are far removed from the frontal margin. The amply ramified intestine begins at or slightly behind the eyes. The pharynx is sit- uated behind the middle of the body, its length amounting to about '/¢ the 390 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON A B Fig. 1. Phagocata holleri. A, Photograph of living specimen, <5.7; B, Outline drawing of living specimen, showing location of eyes, pharynx, and copulatory complex. body length. The copulatory complex appears in the living specimen as a transparent spot in the middle of the postpharyngeal region. Anatomy.—Serial sections of the 4 type-specimens were used in the anal- ysis of the anatomy. The two eyes are situated rather deep below the dorsal epithelium and are of very much reduced size, the diameters of their pigment cups measuring only 13-23 wm. The 2 ovaries, each bearing a conspicuous parovarium, are located below or behind the fourth to sixth pair of lateral branches of the anterior intestinal trunk. The numerous rounded testes (Fig. 2) occupy a broad longitudinal band on either side of the body, extending from behind the ovaries to the region of the pharynx. They are essentially ventral, only very few of them reaching dorsally between the intestinal branches. Many testes are widely connected with the anterior part of the seminal duct or vas deferens (vd) that runs along the medial side of the ventral nerve cord, without forming efferent ductules. The vas deferens expands in the region of the pharynx to form the spermiductal vesicle or false seminal vesicle that proceeds caudally in a tortuous fashion to approach the bulb of the penis. The copulatory apparatus (Fig. 3) is situated some distance behind the pharyngeal pouch. The genital aperture or gonopore (gp) lies about midway between the mouth opening and the tail end of the body. It leads into a small common atrium (ac) that connects dorsally with the duct (bd) of the VOLUME 92, NUMBER 2 391 Fig. 2. Phagocata holleri, photomicrograph of sagittal section through region of testes. i, intestine; t, testis; vd, vas deferens. copulatory bursa and anteriorly with the male atrium (am) surrounding the penial papilla. The atria are lined with a nucleate cuboidal epithelium, some- what thicker in the common atrium than in the male atrium. The penis consists of a moderately muscular bulb (bp) embedded in the mesenchyme and a short, more or less conical papilla (yp) projecting into the male atrium. The large penial lumen (vs) is uniform, not divided into an anterior vesicle and a posterior duct, as is usually the case. It extends as a large cavity from the penial bulb obliquely posteroventrally, tapering toward its opening at the tip of the papilla. In all four specimens the lumen was filled with finger- shaped processes projecting from its wall. The two expanded vasa deferentia (vd) approach the penial bulb from the sides, proceed dorsomedially, and open separately, but close together, into the anterior part of the penial lumen. The copulatory bursa (b) shows no peculiarities. Its outlet, the bursal duct (bd), runs posteriorly above and somewhat to the left of the penis, then curving ventrally, gradually increasing in diameter. It opens into the com- mon genital atrium from the dorsal side. The lining of the entire duct, as well as that of the common atrium, appears to be uniform histologically, so 392 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 0.5mm odc bd ! t | bp vd vS pp Fig. 3. Phagocata holleri, semidiagrammatic view of copulatory apparatus in sagittal sec- tion. ac, common atrium; am, male atrium; b, copulatory bursa; bd, bursal duct; bp, penis bulb; gp, gonopore; odc, common oviduct; pp, penis papilla; vd, vas deferens; vs, penial lumen (seminal vesicle). that it is difficult to pinpoint the boundary between them. There is no dif- ferentiated posterior section of the bursal duct that could be considered a “vagina.” The two oviducts unite in the space between the male atrium and the bursal duct to form a common oviduct (odc) that opens into the male atrium near the transition between the male and common atria. Distribution and ecology.—Five immature specimens of Phagocata hol- leri were collected by Dr. Cato O. Holler, Jr. in Mt. Jefferson Cave, located at an elevation of about 3560 feet in Mt. Jefferson State Park, Ashe County, North Carolina, on 27 May 1978. They were sent to me alive and were placed in a culture maintained at 14°C and fed pieces of beef liver at weekly intervals. At my request, Dr. Holler kindly revisited the cave in September 1978, but found that the seep, in which the original specimens were col- lected, had almost dried up. Two more worms, in poor physiological con- dition, were collected but did not survive the transportation to Washington. The specimens from the original collection, however, matured in the culture and were preserved in August, September, and November 1978. The species appears to be a true troglobite. Taxonomic position.—Among the white species of the genus Phagocata occurring in the Appalachian region, P. holleri shows externally a remark- able similarity to P. angusta Kenk (1977), reported from a cave in West VOLUME 92, NUMBER 2 393 Virginia. Both are characterized by having very slender bodies, truncated heads lacking prominent auricular appendages, the anterior end being some- what more flared in P. holleri than in P. angusta. Both species have ex- tremely small eyes located at a considerable distance from the frontal mar- gin. Anatomically, the two species coincide in having ventral testes confined to the prepharyngeal region. In the structure of the copulatory complex, however, they show several conspicuous differences. The configuration of the penis of P. holleri, with its large villous lumen, is unique among the species of the genus. Other differences concern the histology of the bursal duct. The species is named in honor of its collector, Dr. Cato O. Holler, Jr. of Old Fort, North Carolina. Literature Cited Kenk, R. 1977. Freshwater triclads (Turbellaria) of North America. X. Three new species of Phagocata from the eastern United States.—Proc. Biol. Soc. Washington, 89:645-652. Department of Invertebrate Zoology, National Museum of Natural His- tory, Smithsonian Institution, Washington, D.C. 20560. PROC. BIOL. SOC. WASH. 92(2), 1979, pp. 394-398 NOTES ON TWO SPECIES OF STOMATOPOD CRUSTACEA FROM PHUKET ISLAND, THAILAND Raymond B. Manning Abstract.—Clorida rotundicauda (Miers, 1880) and Cloridopsis benga- lensis (Tiwari and Biswas, 1952) are recorded from mangrove habitats on Phuket Island, Thailand. Miers’ species is shown to be a senior synonym of Squilla choprai Tweedie, 1935, originally described from localities in Malaysia. In 1977 Dawn W. Frith, then with the Phuket Marine Biological Center, Phuket, Thailand, forwarded for identification three specimens of stomato- pods collected during a study of the macrofauna in mangroves at Ao Nam- Bor [07°51'’N, 98°25’E], southeastern Phuket Island, southern Thailand. Two of these specimens were identified with Clorida rotundicauda (Miers, 1880), previously known from one specimen from Formosa, the type-local- ity, and one from Tsimei, near Amoy on mainland China (Schmitt, 1931). The third specimen proved to be Cloridopsis bengalensis (Tiwari and Bis- was, 1952), known from several localities along eastern India (Tiwari and Ghosh, 1975). Further study of the specimens of C. rotundicauda from Phuket had a surprising result: a second species described from mangrove swamps in Malaysia, Clorida choprai (Tweedie, 1935), can be identified with C. ro- tundicauda. Neither C. rotundicauda nor C. bengalensis were included in a recent survey of the stomatopods of Phuket Island, based primarily on species from coral rubble and shale (Dingle, Caldwell and Manning, 1977). Brief accounts of these two species are given below to supplement that account. I thank L. B. Holthuis, Rikksmuseum van Natuurlijke Historie, Leiden, R. W. Ingle, British Museum (Natural History), London, and K. K. Tiwari, Zoological Survey of India, Calcutta, for providing working space and ac- cess to study collections under their care; Dawn Frith, for sending this material for examination; and Lilly King Manning for preparing the illustra- tions. Clorida rotundicauda (Miers, 1880) Fig. 1 Chloridella rotundicauda Miers, 1880:3 [listed], 15, pl. 2, figs. 5, 6—Schmitt, 1931:130, pl. 16, figs. 3-5. VOLUME 92, NUMBER 2 395 Fig. 1. a-g, Clorida rotundicauda, male, 79 mm, Phuket: a, Anterior part of body; b, Eye; c, Claw; d, Lateral processes of fifth, sixth and seventh thoracic somites; e, Sixth abdominal somite, telson and uropod; f, Submedian denticles of telson, ventral view; g, Basal prolongation of uropod. h-i, Male, 51 mm, Tsimei: h, Basal prolongation of uropod; i, Anterior part of carapace and rostral plate. Squilla choprai Tweedie, 1935:49, pl. 1.—Moosa, 1973:147. Material.—Ao Nam-Bor, Phuket Island, Thailand; mangrove mud; D. Frith, leg., zones 3A, 3B, 24 June 1977:2 6, total lengths 76 and 79 mm. Diagnosis.—Eye small, stout, mesial margin flattened. Cornea very small, narrower than stalk, bilobed. Rostral plate triangular, broader than long, lacking carina. Carapace lacking median carina. Anterolateral angles of car- apace occasionally unarmed. Mandibular palp and 3-4 epipods present. Dactylus of claw with 4—6 (usually 6) teeth, proximal very small. Lateral process of fifth thoracic somite bluntly pointed or rounded, unarmed. Pos- terior 3 thoracic and anterior 4 abdominal somites lacking submedian cari- 396 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON nae, latter occasionally faintly indicated on fifth somite. Abdominal carinae spined as follows: submedian 6, intermediate (5)6, lateral 6, marginal 5. Telson denticles 1-3, 5—6, 1. Surface of telson with raised prominences. Submedian teeth of telson with movable apices. Outer margin of uropodal exopod with 3-6 (usually 4) movable spines, distalmost enlarged, spatulate. Inner margin of basal prolongation with 4—7 large fixed spines. Size.—Total lengths of adults 47.5 to 79 mm. Remarks.—The specimens from Ao Nam-Bor were compared with the holotype of C. rotundicauda, a female 70 mm long, Schmitt’s specimen from Tsimei, China, a male 51 mm long, and with accounts and illustrations prepared from syntypes of C. choprai examined in the collections of the Rijksmuseum van Natuurlijke Historie, Leiden (72 , 47.5—58.5 mm) and the Zoological Survey of India, Calcutta (4¢, 20-48 mm and 5°, 22-43 mm). No major differences were observed. The northern specimens and those from Phuket lack the anterolateral spines on the carapace; they are present on all of Tweedie’s specimens examined by me, although he noted (1935:49) that spines were present only in the larger specimens he examined. The smaller specimen from Phuket has a minute denticle at the anterolateral angle of the carapace. Schmitt’s specimen from China differs from the remainder in having a slightly longer rostral plate (Fig. 1/), in having faint submedian carinae on the fifth abdominal somite, in lacking any trace of spines on the intermediate carinae of that somite, and in having a much broader lobe between the spines of the basal prolongation of the uropod (Fig. 14). Otherwise it appears to agree with other specimens in all major respects. It may well represent a distinct species, but this species is so variable I would hesitate to recognize a related species on the basis of a single specimen. One of the specimens from Ao Nam-Bor has but three epipods; the other and Schmitt’s specimen as well, each has four, as do the types of C. choprai. Habitat.—Landward and middle parts of the mangrove forest (zones 3A, B; Frith et. al., 1976:6) at Ao Nam-Bor. Tweedie’s specimens also were taken in mangrove swamps. Distribution.—Indo-West Pacific; records in the literature include: Taiwan: (Miers, 1880).—China: Tsimei [Chi-Mei, 24°30’N, 118°07’E], mainland near Amoy (Schmitt, 1931).—Malaysia: Port Swettenham [03°00'N, 101°24’E], Selangor (Tweedie, 1935).—Singapore: Serangoon Riv- er [Sungai Serangong, 01°24’N, 103°55’E] (Tweedie, 1935; Moosa, 1973). Cloridopsis bengalensis (Tiwari and Biswas, 1952) Squilla bengalensis Tiwari and Biswas, 1952:352, fig. 1b, c. Cloridopsis bengalensis.—Tiwari and Ghosh, 1975:33, fig. 1. VOLUME 92, NUMBER 2 397 ?Chlorida rotundicauda.—Frith, Tantanasiriwong and Bhatia, 1976:14, 19, 35. Material.—Ao Nam-Bor, Phuket Island, Thailand; mangrove mud; D. Frith, leg., zone 2; 24 June 1977:1 6, total length 105.5 mm. Diagnosis.—Eye slender, stalk width about % eye length, inner margin of stalk not expanded, cornea broader than stalk. Rostrum about as long as broad, with median carina anteriorly. Median carina of carapace lacking anterior bifurcation. Mandibular palp and 2 epipods present. Dactylus of claw with 6 teeth. Lateral process of fifth thoracic somite lacking a black spot. Abdominal carinae armed as follows: submedian 6, intermediate S—6, lateral (1-3) 4-6, marginal 1-5. Telson denticles 3—5, 5-6, 1. Apices of sub- median teeth of telson minute, movable. Telson lacking postanal keel. Outer margin of uropod with 6—7 short movable spines. Size.—Tiwari and Ghosh (1975) recorded specimens ranging in size from 56 to 124 mm. Remarks.—The single specimen examined agrees well with the two ac- counts of this species in the literature as well as with figures of a specimen from India prepared by L. K. Manning in 1972. It differs from previous accounts in that the rostrum is as long as broad rather than slightly broader, and that the lateral carinae of the anterior three abdominal somites are unarmed. The carina of the rostrum in this specimen is damaged. Only two species of Cloridopsis from the Indo-West Pacific region are known to have a mandibular palp: C. bengalensis, in which the claw is armed with 6 teeth, and C. terrareginensis Stephenson, 1953, from Austra- lia, in which the claw is armed with 5 teeth. The only other Cloridopsis from the area, C. scorpio (Latreille, 1828), which occurs on mudflats at Ang Sila, lacks the mandibular palp, has an elongate rostral plate, and has a prominent black spot on the curved lateral process of the fifth thoracic somite. Habitat.—In mud at the landward edge of the mangrove forest. Frith et al. (1976:14) noted that this species (as Chlorida rotundicauda) was not found at other zones, and on p. 19 they note that one specimen was taken from a long horizontal burrow about 10 centimeters below the surface in zone 2. Apparently true Clorida rotundicauda subsequently was taken in zone 3 along with additional material of C. bengalensis. Tiwari and Ghosh (1975:37) noted that this species occurred in estuarine habitats in India. Distribution.—Indo-West Pacific; Bay of Bengal, eastern coast of India, from localities in lower West Bengal State in the north to Andhra Pradesh State in the south, and now from Phuket Island, Thailand. It has not been recorded previously from localities outside India. 398 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Literature Cited Dingle, H., R. L. Caldwell, and Raymond B. Manning. 1977. Stomatopods of Phuket Island, Thailand.—Phuket Marine Biological Center, Research Bulletin number 20: 1-20, figures 1-11. Frith, Dawn W., R. Tantanasiriwong, and O. Bhatia. 1976. Zonation of macrofauna on a mangrove shore, Phuket Island.—Phuket Marine Biological Center, Research Bulletin number 10:1-—37, figures 1-3. Miers, E. J. 1880. On the Squillidae.—Annals and Magazine of Natural History, series 5, 5:1— 30, 108-127, plates 1-3. Moosa, M. Kasim. 1973. Type specimens of Crustacea formerly in the National Museum, Singapore.—Journal of the Singapore National Academy of Science 3(2):142—149. Schmitt, Waldo L. 1931. Chinese Stomatopods collected by S. F. Light.—Lingnan Science Journal 8[1929]:127-155, plates 16-19. Tiwari, K. K., and S. Biswas. 1952. On two new species of the genus Squilla Fabr., with notes on other stomatopods in the collections of the Zoological Survey of India.—Rec- ords of the Indian Museum 49(3—4):349-363, figures 1-5. Tiwari, K. K., and H. C. Ghosh. 1975. Redescription of Squilla bengalensis Tiwari and Biswas (Crustacea: Stomatopoda).—Proceedings of the Zoological Society, Calcutta, 26[1973]:33-37, figure 1. Tweedie, M. W. F. 1935. Two new species of Squilla from Malayan waters.—Bulletin of the Raffles Museum 10:45—52, plate 1. - Department of Invertebrate Zoology, National Museum of Natural His- tory, Smithsonian Institution, Washington, DC 20560. PROC. BIOL. SOC. WASH. 92(2), 1979, pp. 399-413 A NEW CRAB FAMILY FROM SHALLOW WATERS OF THE WEST INDIES (CRUSTACEA: DECAPODA: BRACHYURA) Austin B. Williams Abstract.—Mimilambrus wileyi, new genus and new species is described from nearshore waters of Man-of-War Bay, Tobago, West Indies. Mimilam- brus represents a new family that combines characters of the Parthenopidae, Leucosiidae and Calappidae. Its placement, therefore, lies between the Ox- ystomata and Brachygnatha, probably within the Oxyrhyncha. Introduction In April, 1978, participants from the University of Maryland Chesapeake Biological Laboratory conducted a two-week fisheries survey in near-shore and freshwater habitats of Tobago, West Indies, collecting fishes and in- vertebrates. Members of this party were J. D. Hardy, Jr., leader, L. Lubbers III, F. D. Martin, D. Shelton, and M. L. Wiley; the latter asked me be- forehand whether decapod crustaceans would be of interest and was pri- marily responsible for both collecting and transmitting decapods to me after the expedition was completed. Among these specimens, the only brachy- urans were two adults of the extraordinary species described below and named in honor of Dr. Wiley. A third specimen was later transferred from the North Carolina State Museum of Natural History, Raleigh. I am grateful to all members of the University of Maryland party whose efforts made this report possible, especially Dr. Wiley who preserved the specimens for study and Mr. Hardy who initiated the survey, as well as to Mr. H. E. Wood, senior Fisheries Officer, Ministry of Agriculture, Lands and Fisheries, Fisheries Division, Port-of-Spain, Trinidad and Tobago, who was responsible for giving permission to make collections. The manuscript was critically reviewed by F. A. Chace, Jr., D. M. Cohen, B. B. Collette, B. Kensley, R. B. Manning, and M. L. Wiley. Maria Dieguez prepared the illustrations. J. E. Cooper provided the paratype male. Mimilambridae, new family Chelipeds much longer and heavier than other legs; last legs normal in position. Carapace subcircular, barely concealing bases of legs; pointed ros- trum present. Mouth field truncate triangular; external maxillipeds more or less covering buccal cavern, exognath hidden beneath endognath, palp of 400 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON endognath exposed; afferent openings lateral to external maxillipeds and anterior to bases of chelipeds. Antennules folding somewhat obliquely. An- tennae small, peduncles in orbital hiatus. Male openings coxal, female open- ings sternal. The family contains the following genus. Mimilambrus, new genus Carapace subcircular, regions fairly well delineated; numerous forward- trending spines and tubercles tending to arrangement in rows; spined an- terolateral margin ending in strong, suberect, forward-trending lateral spine. Front with small but prominent, depressed rostrum flanked by notches for antennules. Orbits fairly large; eyes retractile, cornea well developed. Third maxilliped with subtriangular merus and broader polygonal ischium, both fringed laterally by dense long setae covering afferent branchial channels; merus with mesial edge grooved for reception of well developed palp, ven- tral edge of groove emarginate. Chelipeds slightly unequal, long, strong, merus and propodus prismatic in cross section, angles densely and strongly spined; fingers extended not diverging from axis of chela, occlusal surfaces with double row of variable teeth incompletely coalesced transversely. Male abdomen with segments 3-5 articulated but not completely mobile; mature female abdomen broad, segments free. Type-species.—Mimilambrus wileyi, new species. Etymology.—From the Latin, *“‘mimus’’ an imitator, and “‘lambrus”’ re- ferring to the old name introduced by W. E. Leach for crabs of the genus Parthenope. Leach gave no derivation for this name. Lambrus is the Latin name of the Lambro River in Lombardy, Italy; many crustacean genera were named for rivers (L. B. Holthuis, personal communication). It is also from Lambrusca, a corruption of Labrusca, a name for wild grapes (Dic- tionnaire des Sciences Naturelle, 1839). Mimilambrus wileyi, new species Figs. 1-5 Description.—Superficially parthenopid-like, with extremely spiny, tu- berculate, granulate and punctate integument; hairs not hooked. Carapace depressed; surface uneven, regions fairly well marked and fairly eroded in irregular pattern more or less corresponding to depressions between raised regions; both eroded and raised areas granulate and punctate. Anterolateral margin with 2 small obtuse spines on subhepatic margin, posterior less than twice size of anterior, followed by spineless interval; 7 marginal spines on branchial margin, first small, second to fifth well formed (second to fourth forward trending), followed by reduced tuberculous sixth spine and line of granules ending in strong, erect, acute, forward-hooked spine at lateral angle 401 VOLUME 92, NUMBER 2 ‘poyeoIpul WW | ‘SUISIeW [e}IGIO pue [eJUOI ‘q ‘poyeoIpUl WU ¢ ‘MOIA [eSIOg ‘B ‘adAjojoy afew ‘Mazi snaquipjUnp ‘| ‘Si 402 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 2. Mimilambrus wileyi, fingers of chelipeds. Female allotype: a, Right, dorsomesial aspect; b, Right, ventrolateral aspect; c, Left, ventrolateral aspect; 5 mm indicated. about twice as long as preceding spines. Strong, elongate, blunt subbran- chial spine above first walking leg (p2). Posterolateral margin straight and granular above base of fourth leg but excavate and emarginate above base of fifth leg. Posterior margin arched ventrad, paralleled by submarginal row of tiny granules. Carapace with median raised tract extending from gastric to posterior margin bearing 5 spines in midline; 1 strong mesogastric preceded by sub- median pair of granular tubercles and these in turn preceded by 3 remote pairs of granular tubercles to either side of midline on proterogastric margin, posterior pair obsolescent; 1 strong urogastric; 2 weak cardiacs flanked by 3 small tubercles at each side; 1 strong, forward-hooked intestinal flanked by pair of tubercles to either side paralleling posterior margin. Branchial regions inflated, with concentrically arched rows of spines and tubercles directed anteromesially from posterolateral margin; 2 low mounds, one above other, at junction of posterior and posterolateral margins on low ridge confluent with cardiac region, lateral to them an arcuate groove; then an ill— defined raised tuberculate area paralleled laterally by row of 5 spines orig- inating above base of fifth leg, last 2 spines acute and hooked forward, penultimate strongest, third spine obtuse or reduced and 2 preceding it re- duced to granular tubercles; lateral to these a row of 4 rather small low spines originating near lateral spine, anteriormost reduced but fairly acute, second and third stronger with accessory granules on slopes, posteriormost low; intercalated between this row of spines and those of anterolateral mar- gin a row of about 3—5 suppressed spines plus granules. Crescentic patch VOLUME 92, NUMBER 2 403 of 3 or 4 granular hepatic tubercles above marginal subhepatic spines, an- teriormost largest, patch confluent with raised orbital region. Front narrow, cut into rostrum consisting of narrow, depressed but pro- jecting triangular tip flanked basally by raised granulate knob at either side, a deep lateral notch, and prominent narrowly triangular inner orbital spine barely exceeding basal rostral knobs and depressed to same level as rostral tip; notch and raised basal knobs provide recesses for obliquely but almost vertically folding antennules. Deeply cut orbits slightly tubular, dorsal mar- gin finely and evenly granulate, single dorsal suture; outer orbital tooth strong, equaling rostrum, flanked by lesser tubercle on upper and lower side, lower one largest; strong infraorbital tooth exceeding rostrum and in turn exceeded by antennal peduncle; latter not filling orbital hiatus. Eyes well developed, subcylindrical stalks of moderate length, retractile, black cornea scarcely dilated. Antennules and antennae short, of about equal length. Mouth field truncate triangular; major articles of third maxillipeds not completely closing mouth field, rimmed by dense fringe of long setae most conspicuous along lateral margins; setae originating as well on sternite III, coxa and basis of chelipeds, and, together with similar setae on sharply defined milled edge of otherwise smooth subbranchial and pterygostomian regions, angling forward to base of infraorbital spine to form filter for hol- lowed afferent branchial channels mesial to milled edge. Marginal notch between pterygostomian and subbranchial regions. Third maxillipeds with ischium and merus punctate and granulate ven- trally; ischium roughly 4-sided, with shallow longitudinal depression; distal corners almost right-angled, proximal corners rounded; lateral margin nearly Straight but oblique and also curved inward proximally; basal margin slightly concave, directed anteromesially; mesial margin nearly straight except an- gled in distal part for lodgment of dactyl, submarginal external line of uni- form fine granules paralleled mesially by stiff submarginal setae overlying row of close-set, almost uniform marginal teeth, teeth and part of setae concealed along lodgment for dactyl; distal margin sinuous, width about 0.6 greatest width of entire article. Merus roughly triangular, reaching level of epistome, slightly longer than broad, ventral surface convex along axis ex- cept shallowly cupped mesially; lateral margin an almost uniform shallow arc; tip rather narrowly rounded; mesial margin split longitudinally; exposed irregular ventromesial part paralleled by submarginal row of nearly uniform close-set granules except where clumped behind jutting proximomesial an- gle; projecting dorsomesial part hidden by palp, margin entire. Distal 3 ar- ticles (palp) of limb smooth and iridescent on exposed surfaces, dense tract of partly concealed setae along mesiodorsal surface ending in long terminal tuft. Carpus transverse in repose, inserted near internal midlength of merus, 404 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 3. Mimilambrus wileyi, mouth field including anterior sternites, bases of chelipeds, etc.; 5 mm indicated. prismatic, strong ridge above insertion of short subcylindrical propodus; dactyl asymmetrically conical; propodocarpal articulation in line with that of ischiomeral articulation. Epistome narrow, partly concealed by long setae on mouthparts, its rather broad, concave, ciliated median notch forming, with associated laciniae of first maxillipeds, a central efferent branchial opening. Chelipeds much longer, stronger and spinier than remaining legs, right crusher slightly larger than left cutter; spines trending forward, tending to alternate large and small along rows, those of dorsal side ciliated and those of outer margin tending to be flattened, especially at tips; merus and pro- podus prismatic in cross section, latter most markedly so; merus separate from basis-ischium. Right cheliped of holotype male: Coxa and basis each with single spine on anterior margin ventrally. Ischium with 5—6 low spines on anteroventral border, penultimate one largest, single anterior spine in line with anterome- sial row of merus; irregular longitudinal row of low tubercles above it. Merus with rows of spines along angles (in order of decreasing average size), along outer (ca. 20 major + smaller intermediate spines), inner (ca. 15 major + VOLUME 92, NUMBER 2 405 Fig. 4. Mimilambrus wileyi, diagram of mouth field including bases of chelipeds, etc., showing branchial channels. Third maxilliped in situ at left showing extensive filter along its lateral side and at base of cheliped; third maxilliped on right turned aside to reveal dorsal aspect of afferent channel; en, Endognath; i, Ischium; m, Merus; ex, Exognath, hidden under endognath. I, Endognath lacinia of first maxilliped. af, Afferent channels. ef, Efferent channels merged in median opening. ch, Base of cheliped. t, Telson. IV, Fourth sternite. smaller intermediates) and mesioventral (ca. 8 + tubercles); dorsal side with row of much smaller spines (ca. 21 + smaller) along length of middle, another row of still smaller spines parallel to inner margin, and tract of scattered obsolescent spines and tubercles adjacent to outer margin; scat- tered tubercles elsewhere. Carpus strongly bent to fall in line with propodus; inner row of tubercles and spines ending in strong, upturned distal spine, smaller internal angle below it spinelike; 1 or 2 small, blunt spines aligned perpendicular to distal margin and lateral to inner spine; central dorsal and intercalated rows of spines diminished to tubercles; spines of external mar- gin (12) more or less increasing in size distally but penultimate largest. Propodus (palm) with row of spines on each angle—outer (15 + smaller), inner (18), and row of small spines (ca. 27) on low dorsal ridge paralleling outer margin, its distalmost at base of dactyl; inner row of spines more 406 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON uniform than others and with 1 or more tubercles on each dorsally; longi- tudinal ventral keel reaching from prominent carpal condyle to tip of fixed finger almost smooth but few granules above it on inner face. Fingers short, extended, not diverging from axis of chela, slender; tips hooked toward each other and crossing, white in color like ventral aspect of chelipeds. Dactyl broad proximally, tapering to point closing inside fixed finger; dorsal rows of spines continued from propodus; inner row with 3 short proximal spines, 1 large erect spine at 4% length followed by 3 small spines, strong obliquely upcurved subdistal spine, and subterminal low tu- bercle; lateral row with 2 unequal proximal spines, second spine matching opposite mesial spine at 4% length, followed by irregularly spaced tubercles and a low spine; low median dorsal ridge with 2 proximal tubercles; occlu- sive surface with double row of 5 somewhat remote teeth coalesced side- by-side into molariform crushers and distributed along proximal % of length, first 3 largest; distal to these a large crushing tooth with flanking anterior cusp followed by smaller pyramidal teeth near tip, proximal pyramidals flanked internally by 5 minute conical teeth. Fixed finger nearly straight but offset from ventral plane of palm; narrower than dactyl and slightly deflect- ed, inner surface very slightly concave but outer surface evenly curved; short subdistal tooth on ventral margin preceding hooked tip; occlusive surface with shorter row of 4 unequal molariform teeth similar to those of dactyl followed by 3 remote pyramidal teeth flanked mesially by shallow slot for reception of opposed largest pyramidal tooth of dactyl; edge of dactylar socket bearing fine granules. Left cheliped comparable to right: Number large spines on meral angles— outer, ca. 19+ smaller intercalary; inner, ca. 15+ tubercles; mesioventral, 10; dorsal side with row of much smaller spines (ca. 18+ smaller) along length of middle; another row of similar spines parallel to inner margin. Carpus essentially as on right. Spines on propodal (palmar) angles—outer, 18+; inner, 20; dorsal row paralleling outer margin, 24; longitudinal ventral keel slightly roughened with obsolescent, forward trending tubercles. Dactyl basically similar to that of right side but narrower; inner dorsal margin with 2 moderate spines and 1 small proximal, enlarged, erect spine at %4 length followed by 2 smaller spines; strong obliquely upcurved subdistal spine and subterminal tubercle beyond it; outer dorsal row with small proximal spine, 2 large nearly equal spines followed by remote tubercle and some granules; low median dorsal ridge with 2 obsolescent spines proximally; occlusive surface with continuous but uneven row of broad, thin shearing teeth run- ning nearly full length of finger, flanked laterally by 6 remote reduced conical teeth in distal half of row, and internally by dense proximal tuft of silky hair, faint axial row of granules along inner surface. Fixed finger somewhat similar to that of right but with inner surface nearly straight, outer margin VOLUME 92, NUMBER 2 407 inflated, finger broadest at proximal % of length; short subdistal conical tooth on ventral margin preceding hooked tip; occlusive surface spooned, row of cutting teeth running along each margin; lateral row arched dorsally in middle, bearing 8 somewhat separated triangular teeth, third to fifth from proximal end of row largest; mesial row straighter and more continuous, composed of about 12 broader irregular teeth. Chelipeds of allotypic female: Similar to male except slightly smaller and usually somewhat fewer spines in rows; lower side of palms more sharply tuberculate; fingers more delicate and with slightly different tooth pattern. Right dactyl with inner dorsal margin proximally to distally bearing 2 mod- erate, 1 large, 4 small, 1 large upcurved spine(s), and a distal tubercle; outer margin bearing 2 large, 2 tiny and 1 small spine(s); middorsal ridge with 2 larger tubercles + accessories proximally; occlusive surface with double row of 5 coalesced crushing teeth, proximal 3 largest, sixth large but more strongly coalesced, seventh small, similar, small eighth and large ninth tri- angularly pyramidal, tenth minute. Fixed finger with subdistal ventral tooth large, acute and exceeding tip of finger; occlusive surface with double row of 5 coalesced crushing teeth but distalmost of these strongly coalesced into triangular pyramid,.remote small tooth beyond it flanked mesially by linear notch and low ridge for reception of large ninth tooth of dacty]l. Left cutter with inner dorsal margin of dactyl proximally to distally bear- ing 1 low secondarily tipped, 2 moderate, | large, 3 small, and 1 large upcurved spine(s) and 1 remote tubercle; middorsal ridge with 2 sharp prox- imal tubercles; outer margin bearing 1 tiny, 2 large spine(s) and 1 remote tubercle; occlusive surface with 11 teeth in row tending to be doubled, 1- 4 single and conical, 9 and 11 flattened in plane of finger axis and with accessory cusps on side adjacent to 10; middle of row flanked laterally by remote dorsal row of about 7 granules or tiny tubercles. Fixed finger with subdistal ventral tooth large curved and acute, exceeding tip of finger; oc- clusal surface spooned; bifurcate slender tooth proximally at edge of dac- tylar fossa in line with lateral row of 9 essentially separate, triangular cut- ting teeth, 5 and 8 flattened in plane of finger axis and with obsolescent accessory cusps on each slope; inner tooth row almost obsolescent, 2 or 3 triangular teeth at point of highest elevation. Walking legs much smaller than chelipeds, shorter than meri of latter; first 2 walking legs of about equal length, third and fourth progressively longer; fourth longest and strongest; meri with row of spines on upper and lower crests; other surfaces of articles smooth; merus of first walking leg shorter than carpus and propodus combined, of last about equal to carpus and propodus combined; dactyls slender, gently curved, tapering to acute corneous tip, slightly flattened, longitudinally ribbed and grooved. Male abdomen with 2 proximal, broad, free segments visible in dorsal 408 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 5. Mimilambrus wileyi. Abdomen and sternites in ventral view; a, Female; b, Holotype male; 5 mm indicated. Gonopods of holotype male in ventral view; c, First; e, Second; 1 mm indicated; d, Tip of first; 0.5 mm indicated. view; ventrally visible part tapering from broad second segment to narrow, subtriangular telson with sinuous sides, its constricted tip reaching nearly to level at which first legs articulate with sternum; abdominal margins dense- ly ciliated. Short first segment slightly arched, filling space between fifth coxae, its somewhat concave dorsal surface molded to posterior marginal VOLUME 92, NUMBER 2 409 contour of carapace but flared laterally to meet basal articles of fifth legs, posterior margin raised into rounded ridge. Second segment drawn into prominent, thin, 3-lobed flange or stop directed posteriorly, broader than first segment and overlapping basis of fifth legs at each side; median lobe narrowly rounded in outline, lateral lobes irregular; dorsal surface concave, smooth, ventral surface similar but with narrower transverse concavity lim- ited to central half. Third segment as broad proximally as first, roughly trapezoidal in outline, its sides narrowing abruptly from rounded proximo- lateral corners to articulation with fourth segment; transverse median ridge broadly confluent with proximolateral corners, but with narrowed central section flanked by transverse concavity anteriorly and posteriorly. Fourth, fifth and sixth segments subquadrate, progressively narrowed toward telson. Third to fifth segments articulated but not completely mobile. Female abdomen (mature) nearly as broad as sternum, composed of 6 free segments (first 2 visible in dorsal view, remainder, including second, visible ventrally); much narrower, subtriangular or hastate telson broader than long, sides concave, proximal margin biconcave, angles rounded but tip most narrowly so, and reaching level of articulation of first legs with sternum; abdominal margins densely ciliated. First segment narrowest and very short, slightly arched, filling space between bases of fifth legs, its some- what concave dorsal surface molded to posterior marginal contour of car- apace but flared laterally to meet basal articles of fifth legs, posterior margin raised into low rounded ridge. Second segment prominent, 3-lobed thin flange or stop directed posteriorly, broader than first segment and overlap- ping basis of fifth legs at each side, median lobe broadly rounded in outline, lateral lobes irregular; dorsal surface concave, smooth, ventral surface sim- ilar but with narrower concavity limited to central half. Third to sixth seg- ments broadly subrectangular, lateral margins of each rounded, broad pos- terior median notch in each filled by articular membrane or anterior median projection of third to fifth; fourth segment broadest; sixth longest, its pos- terolateral margins sloping. Segments 2—5 bearing well developed, biramous pleopods. Circular female openings with crescentic aperture mesially on sixth sternite. Male openings coxal. First pleopods, viewed ventrally, long and slender, reaching to suture between sternites IV and V; relatively stout, narrowly triangular proximal 4 narrowing at slight inflection into longer more slender central 44, elements of each side nearly touching in midline at suture be- tween sternites VI and VII, then gently diverging in distal 4; tip slightly dilated, bluntly pointed and bifid, conspicuously ringed by circlet of sub- apical, retroflexed spinules, longest mesially. Measurements in mm.—Holotype male (H), allotype female (A), and paratype male (P). 410 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Anterior edge merus, External edge Length Width Depth ischiomeral suture- carpus-base of — Length carapace carapace thorax merocarpal condyle dactyl dactyl H 24.3 26.0* 13.0 R 41.5 R 49.9 Ro» ites 28.47 L 41.5 L 47.9 L 10.9 A 24.4 24.8* 13.1 R 34.1 R 43.4 R827 28.27 L 34.1 L 42.0 j BPs) $8) P 25.8 Pap Se 14.0 R 42.0 R_ 50.1 R_ 13.0 30.47 L 41.5 L 48.0 Ue 119227) * Excluding spines; f including spines; R right; L left. Color.—Paratype male in alcohol 9 March 1979. Carapace grayish on gastric and cardiac regions, becoming darker along rows of spines radiating over branchial regions; lighter, flesh colored band around perimeter, broad- est adjacent to anterolateral, orbital and frontal areas, and in area surround- ing cardiac region; spines and tubercles similar except for gray of bran- chial ridge continued on lateral spine. Entire dorsal aspect of carapace and limbs very finely speckled with rust-red; speckles arranged in reticulate pattern on chelipeds; elongate spots of same color along inner surface of each fixed finger except for base and tip; each dactyl with narrower, shorter spot of same color on inner surface, smaller spots at base of penultimate, upturned spine and scattered along dorsal aspect. Material.—USNM 172222, holotype ¢6, USNM 172223, allotype 2, To- bago, West Indies, Man-of-War Bay, gill net over sand at night, depth ca. 10 ft., 14 April 1978. Collected by M. L. Wiley, F. D. Martin et al. USNM 172264, paratype 6, Tobago, Charlottesville, found dead on beach, 16 December 1978. Collected by A. L. Braswell. Transfer from North Carolina State Museum of Natural History. Remarks.—Mimilambrus wileyi has a striking but only superficial resem- blance to crabs of the subfamily Parthenopinae. Spination of the chelipeds is different, the fingers are extended in the longitudinal axis of the hand rather than set at a mesial angle to the palm, although the fixed finger is slightly offset from the ventral margin of the palm, and double rows of teeth on occlusal edges of the fingers are unlike any tooth rows recorded for the parthenopids. The walking legs are much smaller than the chelipeds, as in the Parthenopinae, but the last walking legs are longer than the first and shaped much as in Acanthocarpus alexandri Stimpson (Calappidae) whose meri are smooth rather than spined on upper and lower margins. The carapace is rounded in outline, roughly octagonal rather than trigonal or pentagonal as in many Parthenopinae, but the front bears a short, pointed rostrum (as in some Parthenope, also Heterocrypta and Mesorhoea) and there is a longitudinal, raised middorsal tract extending from the gastric region to the posterior border that bears a few spines. To each side of this VOLUME 92, NUMBER 2 41] tract are raised, roughly concentric tracts bearing more or less developed rows of spines that remotely resemble the faint branchial surface sculpture in Cycloes (also Mursia) (Calappidae), but the forward trend of both these spines and the strongly acuminate, suberect lateral spines, along with the number and arrangement of anterolateral spines, reduces similarity to Cy- cloes or the parthenopids. The frontal region, deeply cut for the antennular sockets and provided with roomy orbits housing well Sec aber retractile eyes, suggests calappid affinities. The uniqueness of the present species becomes most apparent in veniral aspect. The mouthparts which extend forward within a truncately triangular mouth field are immediately reminiscent of the oxystomes as recognized by Rathbun (1937) but modified by Guinot (1966, 1967) to exclude certain groups (Aethrinae Dana 1852; Hepatinae Stimpson 1871) which she allied more closely to the Parthenopidae. The mouthparts of M. wileyi seem sim- ilar to those of the Leucosiidae (Guinot 1966:761, figs. 23-24) but the re- semblance is a superficial one. The similarities are: 1) truncately triangular mouth field; 2) afferent respiratory channels along sides of mouth field with incurrent openings at base of orbito-antennal area; 3) efferent respiratory channels emptying through a common anterior median opening situated be- tween incurrent openings at either side. The differences are: 1) The third maxillipeds do not close tightly together in the midline, but are slightly gaping. 2) The good-sized palp of each third maxilliped (carpus, propodus, dactyl), articulates near the anterointernal angle of the merus in a manner faintly similar to that in the Parthenopidae, and is exposed as in that group, not concealed under the merus as in the Leucosiidae; the roughly triangular merus is excavated and longitudinally grooved mesially for reception of the exposed palp. 3) The exognath of the third maxilliped is completely con- cealed beneath the broad ischium and merus of the endognath and bears a palp, not exposed and palpless, covering the efferent branchial channel as in the Leucosiidae. 4) The basis of the third maxilliped is exposed, but the coxa is almost hidden. 5) Although the afferent branchial channels open anteriorly, lateral to the endognath of each third maxilliped, each is in fact open along the entire lateral side of its ischium and merus as well as in front of the base of each cheliped but covered along this length by a dense filter of overlapping setae fringing the channel margins; the shallow channels are smooth dorsally, each sealed along its mesial side by the concealed exognath of the third maxillipeds and to some extent by exognaths of the second and first maxillipeds. 6) The endognath lacinia of maxilliped 1 is broad and trun- cate distally, forming, with its member of the opposite side, a deflector for water discharging from the efferent channels through an undivided median notch in the epistome. In short, although the external respiratory organi- zation is roughly analogous to that in the Leucosiidae, it is less rigidly confined, has different mesial and ventral boundaries, and seemingly has a 412 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON much more extensive afferent filtration area. But concealment of the ex- ognath of the third maxilliped may also be an extension of the trend seen in Mesorhoea (Parthenopinae) in which that element is visible lateral to but partly tucked under the endognath. The orbits of M. wileyi are more open than those of either the parthen- opids or leucosiids. The orbital hiatus of leucosiids is rather tightly filled by a small, slender antennal peduncle; in M. wileyi this peduncle is relatively larger and more loosely fitted. The orbital hiatus of parthenopids is filled by both the enlarged basal antennular article and restricted, tiny antennal pe- duncle. Although both of these peduncles fold obliquely in all of the forms discussed above, those of M. wileyi are almost vertical. The flattened body, plane, broad, longitudinally elliptical sternum, male abdomen with 3 rather broad proximal and narrowed distal segments, and the fully segmented but rather loosely apronlike mature female abdomen which covers much of the sternum all bear remote resemblance to those features of the Portunidae. The resemblance, however, is tenuous. Seg- ments of the mature female Parthenope abdomen are broad and articulated, but the telson is broad and rounded rather than rather narrow and hastate as in M. wileyi; that of some transitional parthenopid genera discussed be- low is narrower with pointed telson, but tightly fitted to the sternum. Shape of the mature female majid abdomen, while not uniform, tends toward the form of a subcircular purselike receptacle tightly closed against the sternum; there is complete articulation between segments in some species, but ten- dency to emphasized distal segments in many others with either broadening of the rounded telson or suppression of it when the sixth segment is greatly enlarged in a manner paralleling that seen in leucosiids. The male abdomen of M. wileyi is almost completely articulated in segments 3—5, somewhat as in Parthenope, whereas these segments are fused in portunids, calappids and leucosiids. The posterior sternites are wider and larger in Portunidae, but narrower in other groups discussed above, than in M. wileyi. Other similarities in the sterno-abdominal features of these groups seem too re- mote to be of consequence. The male pleopods seem unique, unlike those known among other brachy- urans. ? Guinot (1966), and carcinologists up until around 1900 whose work she reviewed, regarded certain genera as transitional between the sections Ox- ystomata and Oxyrhyncha. Attention was directed (paraphrasing Guinot) to a common narrowness of frontal region and concentration of sensory organs but a differing conformation of buccal cavity and respiratory arrangement, particularly the position of efferent branchial canals. Most of this attention focused on genera (Cryptopodia, Solenolambrus, and especially Meso- rhoea) which somewhat resemble the Oxystomata in arrangement of the buccal area and, rarely (Aethra), in position of chelipeds, but not in position VOLUME 92 NUMBER 2 413 of antennular and antennal peduncles. Rathbun (1925, 1937) accepted the ideas of Alcock and included these genera in the Parthenopidae (Guinot 1966), as did Garth (1958). Guinot (1967) then extended the transitional status to genera of Oxy- stomes which could be considered as transitional between these two groups (Aethra [again], Osachila, Actaeomorpha, Hepatella and Hepatus). She set up another set of comparisons among these intermediate crabs. Those with either oxystomous or brachygnathous buccal frame were considered par- thenopid in appearance if antennules and antennae fold longitudinally or slightly obliquely, male abdominal segments are distinct, sternum is nar- rowed posteriorly, basis-ischium of cheliped is separate from merus, and exognathal flagellum of the third maxilliped is present. Those with oxysto- mous buccal frame were not considered parthenopid in appearance if an- tennules and antennae are strongly folded obliquely, male abdominal seg- ments 3-5 are fused, sternum is broadened posteriorly, basis-ischium of chelipeds is fused with merus, and exognathal flagellum of the third maxil- liped is reduced or absent. After some collateral comparisons, Guinot pro- visionally proposed uniting the genera Aethra, Osachila, Hepatus, Hepa- tella and Actaeomorpha in the same taxonomic unit, the Aethrinae Dana 1852 which she resurrected and placed in the Parthenopidae near genera such as Cryptopodia. Sakai (1976) adopted this placement. From another point of view, Glaessner (1960:46) considered the parthen- opids with their pointed rostrum, prominent mesogastric-cardiac ridge and elongate [broadened?] cephalothorax to be much closer to the Oxystomata than to the Brachyrhyncha, although in his view their mouthparts are much more advanced [mostly brachygnathous]. According to Glaessner (1969) the Oxystomata first appear in the Lower Cretaceous, but details of their sub- division are obscure. The Leucosiidae, probably derived from the Calap- pidae, do not appear until the Cenozoic. Glaessner regarded origin of the Oxyrhyncha also as obscure, for at their first appearance in the Eocene the families Majidae and Parthenopidae are well differentiated. The Oxystomata have been suggested as their ancestors. Mimilambrus wileyi occupies a position intermediate among these fami- lies. Which of its characters represent convergent specializations and which are fundamentally transitional is unknown. It is perhaps more parthenopid than calappoid. Literature Cited Dana, J. D. 1852. Crustacea. Jn United States Exploring Expedition, during the years 1838, 1839, 1840, 1841, 1842. Under the command of Charles Wilkes U.S.N. 13(1):685 pp. C. Sherman, Philadelphia. Garth, J. S. 1958. Brachyura of the Pacific Coast of America, Oxyrhyncha.—Allan Hancock Pacific Expeditions 21(1):1-xii, 1-499. 414 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Glaessner, M. F. 1960. The fossil decapod Crustacea of New Zealand and the evolution of the order Decapoda.—New Zealand Department of Scientific and Industrial Research, New Zealand Geological Survey. Paleontological Bulletin 31:5—79. . 1969. Decapoda.—iIn R. C. Moore, ed. Treatise on invertebrate paleontology, Pt. R, Arthropoda 4, Vol. 2, pp. R399-533, R626—628, University of Kansas and Geological Society of America, Inc. Guinot, D. 1966. Recherches préliminaires sur les groupements naturels chez les Crustacés Décapodes Brachyoures. I. Les affinités des genres Aethra, Osachila, Hepatus, He- patella et Actaeomorpha.—Bulletin du Muséum National d’Histoire Naturelle, Ser. 2, 38(5):744-762. 1967. Ibid. I. Les Affinités des genres Aethra, Osachila, Hepatus, Hepatella et Ac- taeomorpha (suite et fin).—Op. cit. 38(6) [for 1966]:828-845. Rathbun, M. J. 1925. The spider crabs of America.—United States National Museum Bulletin 129:i-xx, 1-613, 283 pls. . 1937. The oxystomatous and allied crabs of America.—United States National Mu- seum Bulletin 166:i—vi, 1-278, 86 pls. Sakai, T. 1976. Crabs of Japan and the adjacent seas.—Kodansha Ltd., Tokyo, 773 pp. [English text]. Stimpson, W. 1871. Preliminary report on the Crustacea dredged in the Gulf Stream in the Straits of Florida by L. F. de Pourtales, Assist. U.S. Coast Survey. Part I. Brachyura.— Bulletin of the Museum of Comparative Zoology, Harvard College 2(2):109—160. Systematics Laboratory, National Marine Fisheries Service, National Museum of Natural History, Washington, D.C. 20560. a 4 i SAAS, oi tote Ua ih j pt Wgoatca. iit Ye iy "iy ie The ere cee ae 4 Aietitehs rapes LWA! ita Realy My) ‘pont SNe itl) eS | ye; tage % tiara es aa toogehes TA inks, TEMES gi 0 : AY ; er ce macisseiasii cat ai. Re aes see Pry regs. ey * leu oie as nea Nd, ue agaiems! pa I kN ! PEE CERAM WAR Peau Bd CR ARS Ne ay ey eka ae vive fir Ree: Bla PR. SN ST a i Henan Me Joined ee ey | Lape ea ae PRD ait eae es Aged ae ace ghee teap Lie ars ak Wists ie oA oer ie = ere. Geren o.% aa ee on al § hee, pal tae a f oi % 2 Deh, ' i : i & Gh , # i INFORMATION FOR CONTRIBUTORS Content.—The Proceedings of the Biological Society of Washington contains papers bearing on systematics in the biological sciences (both botany and zoology, including paleontology), and notices of business transacted at meetings of the Society. Except at the direction of the Council, only manuscripts by Society members will be accepted. Papers will be published in English, except for Latin diagnosis/description of plant taxa which should not be duplicated by an English translation, or summary in an alternate language when appropriate. Submission of manuscripts.—Manuscripts should be sent to the Editor, Proceedings of the Biological Society of Washington, National Museum of Natural History, Washington, D.C. 20560. Review.—One of the Society’s aims is to give its members an opportunity for prompt pub- lication of their shorter contributions. Manuscripts will be reviewed in order of receipt by a board of associate editors and appropriate referees. The Proceedings are issued four times a year. Presentation.—Clarity of presentation and requirements of taxonomic and nomenclatural procedures necessitate reasonable consistency in the organization of papers. Authors should follow recent issues of the Proceedings as models, including an abstract. Telegraphic style is recommended as the most economical of space for descriptions. Synonymy of abbreviated style (author, date, page) with full citations only in Literature Cited is also recommended. The establishment of new taxa must conform with the requirements of the appropriate in- ternational codes of nomenclature. Authors are expected to be familiar with these codes and to comply with them. New species-group accounts must designate a type-specimen deposited in an institutional collection. The sequence of material should be: Title, Author(s), Address(es), Abstract, Text, Literature Cited, Appendix, Tables (each table numbered with an Arabic numeral and heading provided), _ List of Figures (entire figure legends), Figures (each numbered and identified). Type manuscripts double-spaced throughout (including tables, legends, and footnotes) on - one side of paper measuring approximately 812 x 11 inches, leaving margins of at least one Se aT inch all around. Submit a facsimile with the original and retain an author’s copy. Number pages _ consecutively at the top. One manuscript page = approximately 2 of a printed page. Underline singly scientific names of genera and lower categories; leave other indications to _ the editors. Figures and tables, with their legends and headings, should usually be self explanatory, not _ requiring reference to the text. Indicate their approximate placement by a pencil mark in the margin of the manuscript. Illustrations should be planned in proportions that will efficiently use space on the type bed of the Proceedings (11.8 x 18 cm). Legends require 4 mm of column length per line. Proofs.—Galley proof will be submitted to authors for correction and approval. Changes _ other than printer’s or editor’s errors may be charged to authors. Reprint orders will be taken _ with returned proof. CONTENTS New records of the species of Glyphocrangon in the northeastern Pacific Ocean (Caridea: Glyphocrangonidae) Mary K. Wicksten The crayfishes of New England Denton W. Crocker The marine isopod crustaceans of the Gulf of California II. Idoteidae: new genus and | species, range extensions, and comments on evolution and taxonomy within the family Richard C. Brusca and Barry R. Wallerstein A new species of Leiocephalus (Reptilia: Iguanidae) from Hispaniola Albert Schwartz A new species of tree squirrel (Sundasciurus) from Palawan Island, Philippines (Mammalia: Sciuridae) Lawrence R. Heaney New species of Nearctic Hydroporus (Coleoptera: Dytiscidae) James F. Matta and G. William Wolfe A new species of Ctenocheles (Crustacea: Decapoda: Thalassinidea) from the north- western Gulf of Mexico Nancy N. Rabalais Studies on decapod Crustacea from the Indian River’ region of Florida XIII. Larval development under laboratory conditions of the spider crab Mithrax forceps (A. Milne Edwards, 1875) (Brachyura: Majidae) Kim A. Wilson, Liberta E. Scotto, and Robert H. Gore Mesosignum antarcticum, new species, the first record of the genus from the deep sea south of the Antarctic Convergence (Isopoda: Janirioidea) George A. Schultz A new water scavenger beetle from Mexico (Coleoptera: Hydrophilidae) : Paul J. Spangler and Silvia Santiago-de Bueno Juvenile morphology of the rare burrowing mud shrimp Naushonia crangonoides Kingsley, with a review of the genus Naushonia (Decapoda: Thalassinidea: Laome- diidae) Joseph W. Goy and Anthony J. Provenzano, Jr. Octopus rapanui, new species, from Easter Island (Cephalopoda: Octopoda) Gilbert L. Voss Revision of American species of the marine amphipod genus Paraphoxus (Gammaridea: Phoxocephalidae) J. Laurens Barnard Nannosquilla vasquezi, a new stomatopod crustacean from the Atlantic coast of Panama Raymond B. Manning Redescription of Bruunilla natalensis Hartman eae Polynoidae), originally referred to Fauveliopsidae Marian H. Pettibone Freshwater triclads (Turbellaria) of North America. XI. Phagocata holleri, new spe- cies, from a cave in North Carolina Roman Kenk Notes on two species of stomatopod Crustacea from Phuket Island, Thailand Raymond B. Manning A new crab family from shallow waters of the West Indies (Crustacea: Decapoda: Brachyura) Austin B. Williams 217 225 255) 1 272 280 287 294 307 328 333 339 360 368 380 384 389 394 399 Proceedings of the BIOLOGICAL SOCIETY of WASHINGTON olume 92 18 October 1979 Number 3 sect iain ' a tf —_ Se a THE BIOLOGICAL SOCIETY OF WASHINGTON 1978-1979 Officers President: Oliver S. Flint, Jr. Secretary: Michael A. Bogan Vice President: Richard Banks Treasurer: David L. Pawson Elected Council Arthur H. Clarke W. Duane Hope Robert H. Gibbs, Jr. Catherine J. Kerby Austin B. Williams PROCEEDINGS Editor: C. W. Hart, Jr. Associate Editors Classical Languages: George C. Steyskal Invertebrates: Thomas E. Bowman Plants: David B. Lellinger Vertebrates: Richard Banks Insects: Robert D. Gordon Membership in the Society is open to anyone who wishes to join. There are no prerequisites. Annual dues of $7.00 include subscription to the Proceedings of the Biological Society of Washington. Correspondence concerning membership should be addressed to the Treasurer, Biological Society of Washington, National Museum of Natural History, Smithsonian Insti- tution, Washington, D.C. 20560. The Proceedings of the Biological Society of Washington is issued quarterly in February, May, August and November. Manuscripts, corrected proofs, editorial questions should be sent to the Editor, Biological Society of Washington, National Museum of Natural History, Smithsonian Institution, Wash- ington, D.C. 20560. 7 Known office of publication: National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560. Printed for the Society by Allen Press, Inc., Lawrence, Kansas 66044 Second class postage paid at Washington, D.C., and additional mailing office. PROC. BIOL. SOC. WASH. 92(3), 1979, pp. 415-465 LUMBRINERIDAE, ARABELLIDAE, AND DORVILLEIDAE (POLYCHAETA), PRINCIPALLY FROM FLORIDA, WITH DESCRIPTIONS OF SIX NEW SPECIES Thomas H. Perkins Abstract.—The genus Lumbrinerides is summarized. Lumbrinerides jo- nesi, Lumbrineris ernesti, Lumbrineris verrilli, Protodorvillea bifida, and Schistomeringos pectinata, n. spp., from Florida, Lumbrinerides dayi, n. sp., from North Carolina, and Lumbrineris crosslandi, n. sp., from the Red Sea, are described. Lumbrineris dubeni (Kinberg), L. tenuis (Verrill), L. hebes (Verrill), L. testudinum (Augener), Arabella mutans (Chamberlin), A. multidentata (Ehlers), and A. maculosa Verrill are redescribed. Supple- mental descriptions are provided for Dorvillea sociabilis (Webster) and Pro- todorvillea kefersteini (McIntosh) from Florida. Introduction This report is based primarily on specimens collected during an environ- mental baseline study of nearshore marine fauna near the Florida Power and Light Co., Inc., nuclear power plant at Hutchinson Island, St. Lucie County, southeastern Florida. Some specimens from other areas were ex- amined to clarify taxonomic problems and are also included. A complete report on the Polychaeta of the Hutchinson Island study is in preparation. The study area was characterized and methods of collection were de- scribed by Gallagher and Hollinger (1977). Sediments were described by Gallagher (1977), and other aspects of the physical and chemical environ- ment were reported by Worth and Hollinger (1977). Brief descriptions of benthic sampling stations and methods are given below. Station I (27°21.3’N, 80°14.1’W): seaward margin of beach terrace, about 0.5 km offshore; mean depth about 8.4 m; gray, hardpacked, moderately well sorted, very fine to fine quartose sands. Station II (27°21.6'N, 80°13.2’W): shallow trough ENE of Station I, ap- proximately midway between beach terrace and Pierce Shoal, an offshore bar; mean depth about 11.2 m; poorly sorted, coarse to very coarse calcar- eous sands, with some shell fragments in granule class and minute amounts of silts and clays. Station HII (27°22.0'N, 80°12.4’W): Pierce Shoal, 3 km ENE of Station I; 416 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON mean depth about 7.1 m; well sorted, clean, medium calcareous sands con- taining less than 2% large shell particles. Station IV (27°20.7'N, 80°12.8’W): 1.6 km SSE of Station II in shallow trough about 0.6 km W of southern tip of Pierce Shoal; mean depth about 10.9 m; poorly sorted calcareous sands, predominantly in coarse category; large shell particles accounting for less than 20% sediment weight, minute amounts of fine sands always present. Station V (27°22.9’'N, 80°13.9’W): 2.2 km NNW of Station II; mean depth about 10.9 m; poorly sorted, coarse to very coarse calcareous sands; gran- ule-sized particles consistently more than 25% sediment weight, fine to very fine quartz particles comprising 3—11% sediment weight, mud (clay) fraction less than 1% total weight always present. Samples were taken bimonthly for two years from September 1971 through July 1973 using a Shipek grab and consisted of five replicates cov- ering a total bottom area of about 0.20 m?. Lighter specimens were elutriated on a 0.71 mm screen after return to shore. Remaining sediment was also screened, and specimens were removed under a dissecting microscope. These procedures resulted in collection of many small specimens. Speci- mens were sorted to phyla and transmitted to the Florida Department of Natural Resources Marine Research Laboratory at St. Petersburg for iden- tification. Types and additional specimens available for study are deposited in the following institutions: Allan Hancock Foundation, University of Southern California (AHF); British Museum (Natural History) (BMNH); Florida De- partment of Natural Resources Marine Research Laboratory (FSBC J); Museum of Comparative Zoology, Harvard University (MCZ); Naturhis- toriska Riksmuseet, Stockholm (NRS); National Museum of Natural His- tory, Smithsonian Institution (USNM); Peabody Museum of Natural His- tory, Yale University (YPM); Smithsonian Institution, Ft. Pierce, Florida (SIFP); Zoologisches Museum, Berlin (ZMB); and Zoologisches Museum, Hamburg (ZMH). Maxillae I-V are abbreviated to M I-V in descriptions. Family Lumbrineridae Malmgren, 1867 Genus Lumbrinerides Orensanz, 1973 Type-species.—Lumbrinerides gesae Orensanz, 1973, by original desig- nation. Diagnosis.—Prostomium long, distally pointed. Peristomium divided into 2 short rings. Anterior parapodia very small, almost vestigial. Setae includ- ing simple bidentate hooded hooks and simple limbate setae. Pygidium ter- minating in flattened sucker-like lobe, without anal cirri. Branchiae absent. Mandibles with posterior rami completely fused into a median appendage VOLUME 92, NUMBER 3 417 and distally contiguous. Maxillary supports relatively large. M I broad, with inner borders sinuous or dentate. M II with imperfectly dentate borders or with short, blunt teeth. M III unidentate. M IV in form of more or less rounded adentate plate [after Orensanz, 1973b]. Remarks.—Orensanz (1973b) referred six species to Lumbrinerides; all but the poorly known L. nasuta (Verrill, 1900) from Bermuda are summa- rized, and five additional species are added below, including two new com- binations, two new species and one indeterminable to species. Features of some added species amend the variation of characters known for the genus. Maxillae I of two added species are adentate; mandibles of one are distinctly divided posteriorly; and two are reported to have only one visible peristo- mial segment. Specific diagnostic characters among the species are presently difficult to define, consisting principally of size, number of visible peristo- mial segments, range of appearance of hooded hooks and details of mouth- parts. As aresult, some biological species are apparently clearly distinguish- able from others only on a zoogeographic basis. The original description of L. nasuta (Verrill, 1900:651) stated that uncini are “‘terminated by two small strongly incurved apical hooks, with a large, stout blunt ventral hook,’’ a character indicating the species should be as- signed to Lumbrineriopsis Orensanz, 1973. KEY TO SPECIES OF LUMBRINERIDES IPR OneoVASIDIC PEMSLOMIAlpSeOMENl Livers: aaaGe- leic enc cet ce Z SEV OvISIDIC PelistOmial SCOMEMIS!.. Ske .\. 0. 4s)a- nab he eos Ge we = 3 2. M I with 1 rounded accessory tooth; M II with 3 rounded teeth; jateral supports absent or reduced) 3.) --o.G.. 2. a. ee L. carpinei — MI with 1-2 very small teeth; M II with 2-3 small pointed teeth; luteral supporis long, prominent... 0.2222 425....).. L. crassicephala Bee lvithoumaccessony tect el? . sti aki fic. Seimeiiwen « ATES 4 SE Malandthbaccessony ect haga: ue: fe. be bat Ol. G eed ce Side ee 5 4. Mandibles with small posterior notch; M II with 3 molar-like teeth sy MT MOR es chs AS Esl gins ay a awe Nic teria hhes’ wae Ls JONES. Na SP- — Mandibles widely divided posteriorly; M II with 3 rounded teeth .. eR es Sloe be ee RNS oA Skene coe es oe bus ties Lumbrinerides sp. See Lawati Mp ACCESSORY. LOOT «.cpsityecs. amie bya tml veep tana oo Ie 6 Selenite sACCESSONVAACOUNC. bay sjays lng akys omeste a 2 Siete. AAA yes + 6 n.5 7 6. Accessory tooth of M I as large as distal tooth; hooks beginning in MM ICIOM MOO ested an ce Ih.bs Soke ercsep teat tsps ak Aches RAS SE re L. acuta — Accessory tooth small; hooks beginning on setiger 1 ..... L. aberrans Te ACeeSsoryatecsn ol Mull NEAL AP. oq «ceed st. AAs Be der L. platypygos SS PACCESSOMNtCetm Ol MLL sean MNCS iy. 8428 Dect cteasld sreeeperers |e cers 8 418 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 1. Lumbrinerides acuta (syntype USNM 12895): a, Maxillae. Lumbrinerides aberrans (holotype BMNH ZB 1963:1:86): b, Maxillae. Lumbrinerides dayi: c, Maxillae of juvenile; d, Mandibles, ventral view; e, Maxillae of large specimen. 8. Accessory teeth of M I small; mandibles 3 times longer than broad SR 8h 2 OR EE MAb BER TTR A Oa L. acutiformis — Accessory teeth of M I large, as broad as tip; mandibles 2.2—2.3 times longer than broad VOLUME 92, NUMBER 3 419 9. M II with irregular subequal teeth; mandibles posteriorly tapered, POMC RMN ee mae Ren ec a ca ee cs thea ties cee ars L. gesae — Basal tooth of M II twice as broad as others; sides of mandibles parallel posteriorly, with small posterior notch ........ EV dayinn. so. Lumbrinerides acuta (Verrill, 1875) Fig. la Lumbriconereis acuta Verrill, 1875:39, pl. 3, fig. 5. Lumbrineris acuta.—Hartman, 1942a:114, figs. 10a—d. Lumbrinerides acuta.—Orensanz, 1973b:371.—Gardiner, 1976:203, figs. 26i-1 [in part, New England specimens]. Material examined.—RHODE ISLAND: Off Block Island, 26 m, August 1874, U.S. Fish Comm.; syntype (USNM 13392), 4 syntypes (USNM 12895). Diagnosis. —Two apodus peristomial segments; hooded hooks beginning in middle segments; maxillae delicate, symmetrical: M I (2 + 2) with large accessory tooth near blunt tip subequal to distal tooth; M II (3 + 3) long, with 3 rounded, intermeshing teeth; lateral supports beginning at outer bor- ders of M I, continuing to bases of M III; mandibles about as long as max- illary apparatus, anterior ends flared, posterior ends entire (Hartman, 1942a). Lumbrinerides. aberrans (Day, 1963) Fig. 1b Lumbrineris aberrans Day, 1963:411, figs. 8a—f; 1967:439, figs. 17.17a—e.— Ramos, 1976:108. Lumbrinerides aberrans.—Orensanz, 1973b:373. Material examined.—SOUTH AFRICA: Cape Province, 33°58.8’S, 25°42.2'E, 26 m; holotype (BMNH ZB 1963:1:86). Diagnosis.—Two apodus peristomial segments; hooded hooks beginning on setiger 1 (Ramos, 1976); M I with accessory tooth near blunt tip, much smaller than primary tooth; M II relatively long, with 3 rounded, inter- meshing teeth, anterior tooth half diameter of others; lateral supports be- ginning at about middle of outer edges of M I, continuing to bases of M III; mandibles with posterior ends completely united (Day, 1967). Lumbrinerides crassicephala (Hartman, 1965) Lumbrineris crassicephala Hartman, 1965:117, pl. 20, figs. c-f.—Ramos, 1976:109. Lumbrinerides crassicephala.—Orensanz, 1973b:373. 420 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Type-locality.—Bermuda, in 1000 m. Diagnosis.—One apodus peristomial segment; hooded hooks beginning on setiger 6; M I with acute tip and 1-2 very small accessory teeth; M II short, with 2—3 small, pointed teeth; lateral supports very long, beginning at posterolateral edges of M I, continuing to bases of M III; mandibles nearly as long as maxillary apparatus, 2.4 times longer than broad, poste- riorly pointed [after Hartman, 1965]. Lumbrinerides platypygos (Fauchald, 1970) Lumbrineris platypygos Fauchald, 1970:106, pl. 18, figs. a—d. Lumbrinerides platypygos.—Orensanz, 1973b:373. Type-locality.—Baja California, in shallow subtidal depths. Diagnosis.—Two apodus peristomial segments; hooded hooks beginning on setigers 7-8; maxillae long, delicate; M I with 2 accessory teeth near acute tip; M II of medium size, with 3 rounded, subequal teeth; presence of lateral supports not determined [after Fauchald, 1970]. Lumbrinerides acutiformis (Gallardo, 1968), new combination Lumbrineris acutiformis Gallardo, 1968:82, pl. 29, figs. 2—-8.—Ramos, 1976: 107. Type-locality.—Nha Trang Bay, Viet Nam, in shallow depths. Diagnosis.—Two apodus peristomial segments; hooded hooks beginning on setiger 1 (setigers 1 or 4 fide Ramos, 1976:107); M I with blunt tip and 2 very small accessory teeth near middle; M II with 3 small, rounded teeth, basal one perhaps smaller than others; lateral supports between middle of outer edges of M I and bases of M III; mandibles almost as long as maxillary apparatus, 3 times longer than broad, sides parallel posteriorly, with round- ed tip [after Gallardo, 1968]. Lumbrinerides gesae Orensanz, 1973 Lumbrinerides gesae Orensanz, 1973b:373, pl. 12. Type-locality.—Off Mar del Plata, Province of Buenos Aires, Argentina, in 13 m. Diagnosis.—Two apodus peristomial segments; hooded hooks beginning on setiger 1; M I with blunt tip and 2 accessory teeth near middle, stout, as broad or broader than primary tooth; M II with 3 rounded teeth (irregularly and imperfectly toothed fide Orensanz, 1973b); triangular supports on bases of M III but lateral supports not determined; mandibles about as long as maxillary apparatus, 2.3 times longer than broad, tapered posteriorly, with pointed tip [after Orensanz, 1973b]. VOLUME 92, NUMBER 3 421 Lumbrinerides carpinei (Ramos, 1976), new combination Lumbrineris carpinei Ramos, 1976:109, figs. 5, 6. Localities.—Catalonian Spain and off Monaco in the Mediterranean Sea, in 290-600 m. Type-locality.—Bathyl ‘*‘Etage’’ of the Mediterranean Sea. Diagnosis.—One long apodus peristomial segment; hooded hooks begin- ning on setiger 1; maxillae delicate; M I each with blunt tip and single accessory tooth, near tip on left and near middle on right; M II long, with 3 rounded teeth; lateral supports between M I and M III absent, possibly overlooked; mandibles not visible [after Ramos, 1976]. Lumbrinerides sp. Lumbrineris acuta.—Ramos, 1976:105, figs. 1-3 [not Lumbriconereis acuta Verrill, 1875]. Locality.—Golfo de Rosas, Spain, in shallow depths. Diagnosis.—Two apodus peristomial segments; hooded hooks beginning on setiger 16; maxillae delicate; M I with blunt tip, without accessory teeth; M I of medium length, with 3 rounded teeth; lateral supports between middle of outer edges of M I and bases of M III; mandibles about as long as maxillary apparatus, 3.1 times longer than broad, widely divided poste- riorly. Remarks.—This is apparently an unnamed species, but the wide posterior division of the mandibles may be an artifact of the dissection process. Lumbrinerides dayi, new species Figs. lc—e Lumbrineris aberrans.—Day, 1973:59 [not Day, 1963]. Lumbrinerides acuta.—Gardiner, 1976:203, figs. 26f—h [in part, North Car- olina specimens] [not Lumbriconereis acuta Verrill, 1875]. Material examined.—NORTH CAROLINA: off Beaufort, 34°34'N, 78°25'W, 20 m, sand and broken shell, J. H. Day, col.; holotype (USNM 51145), 17 paratypes (USNM 55598; FSBC I 18478).—FLORIDA: off Pan- ama City, 29°48’N, 86°09’ W, 47 m, coarse carbonate sand, BLM Sta. 2531; R/V Colombus Iselin, Nov. 1977; 1 paratype (USNM 55875).—PUERTO RICO: off Barceloneta (EPA Oceanogr. Study, B. S. Mayo, col.) 18°29'51"N, 66°33’W, 27 m; 7 Aug. 1974; 3 paratypes (USNM 52186).— 18°29'45’N, 66°32'09”W, 30 m; 11 Sept. 1974; 1 paratype (USNM 52187).— 18°29'49"N, 66°32'43”W, 28 m; 12 Sept. 1974; 1 paratype (USNM 52188).— 18°29'51"N, 66°33’ W, 23-28 m; 12 Sept. 1974; 5 paratypes (USNM 52189).— 422 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 18°29’41"N, 66°31'35”W, 34 m; 8 Nov. 1974; 1 paratype (USNM 52190).— 18°29'S51"N, 66°33'W, 28 m; 9 Nov. 1974; 4 paratypes (USNM 52191; FSBC I 18962).—18°29'53"N, 66°33'17"W, 27 m; 10 Nov. 1974; 6 paratypes (USNM 52192).—18°29'51"N, 66°33'W, 23-28 m; 6-8 Aug. 1974; 8 paratypes (USNM 52209). Description.—Colorless in alcohol. All specimens from type-locality pos- teriorly incomplete; longest anterior piece 40 mm long, 0.84 mm wide, 80 segments; anterior piece of 13 setigers, 1.16 mm wide at about setiger 10. Specimen from Gulf of Mexico (USNM 55875) 42 mm long, about 0.8 mm wide, 70 segments. Seven complete specimens from Puerto Rico from 12 mm long, 0.25 mm wide, 75 segments to 33 mm long, 0.5 mm wide, 190 segments. Prostomium 1.2 to more than 3 times longer than wide depending upon state of contraction; tip either acutely pointed (relaxed) or acuminate (contracted); eyes absent. Peristomium with 2 equal apodus segments, to- gether equal in length to setiger |. Parapodia of first 6 segments small, with rounded setal lobes and small, rounded postsetal lobes, gradually increasing in length posteriorly from about setiger 7, fully developed by setiger 10-12; setal lobes short, blunt, conical, slightly longer than wide, about !/,,) as long as body width; postsetal lobes rounded, about as long as setal lobes. An- terior parapodia with 3—4 broadly limbate setae, usually single in posterior parapodia. Simple bidentate hooded hooks beginning on setigers 1—6, con- tinuing to posterior end, 2 hooks at about setiger 10, usually 3 hooks in posterior parapodia. Pygidium as usual in genus, flattened, rounded, about as wide as long, with dorsal anus. Mandibles (Fig. 1d) light to dark brown, 2.3 times longer than broad, delicate, widely flared anteriorly, posteriorly long, narrow, sides almost parallel, about twice as thick as wide, fused, with small posterior notch; length about equal to maxillary apparatus. Maxillary carriers broadly tri- angular, with conspicuously constricted lateral borders much shorter than middle ones, much darker than rest of maxillae. M I lateral borders much longer than medial ones, tips blunt, with 2 rounded accessory teeth near middle, widely separated from tips. M II stout, with 3 rounded, intermeshing teeth, basal one about twice as broad as others. M III relatively stout, with broad, posterior borders. M IV thin, elongate-oval, with narrow posterior borders. Lateral accessory supports from middle of M I to bases of M III; triangular supports at bases of M III. Remarks.—Day (1973) indicated that the number of accessory teeth on Maxilla I might vary among members of L. aberrans, but this was not the case for all of his North Carolina specimens and others I examined; juveniles have the same number and arrangement of teeth on Maxillae I and II as adults (Figs. 1c, e). Hooded hooks first appear on North Carolina specimens from setigers 3—6, while on all other specimens, including a relatively large VOLUME 92, NUMBER 3 423 one from the Gulf of Mexico, hooks begin on the first setiger. The range of first appearance of hooded hooks on anterior setigers may depend on spec- imen size, with hooks first appearing on more posterior segments of larger specimens, or the range may vary between populations from different geo- graphic areas. Hooks may also be broken off anterior ends of dissected specimens. However, mouthpart configurations are constant among all specimens of L. dayi, as well as among specimens of other Lumbrinerides species I have examined. Lumbrinerides dayi is similar to L. acutiformis, L. platypygos and L. gesae and separable from the latter three species as indicated in the key. Additionally, mandibles of L. gesae are posteriorly entire, and, according to Fauchald (1970), postsetal lobes of L. platypygos are always shorter than setal lobes. Etymology.—The species is named in honor of Professor J. H. Day, who collected and first reported the specimens from North Carolina. Lumbrinerides jonesi, new species Fig. 2 Material examined.—FLORIDA: Hutchinson Island, Sta. II, 27°21.6'N, 80°13.2'W, about 11 m, coarse calcareous sand; holotype (R. Gallagher, col., Jan. 1972; USNM 57435); 26 paratypes (USNM 57437; AHF POLY 1268; ZMH _ P-15535-—15537; FSBC I 20434—20444).—Sta. IV, 27°20.7’N, 80°12.8’W, about 11 m, coarse calcareous sand; 9 paratypes (USNM 57436; FSBC I 20445—20450).—Sta. V, 27°22.9’N, 80°13.9’W, about 11 m, coarse calcareous sand; | paratype (AHF POLY 1267). Description.—Generally colorless or pink in alcohol; thoracic region oc- casionally indistinctly rose-colored. Length up to 50 mm, width 1.0 mm excluding parapodia. Prostomium acutely pointed, usually more than twice longer than wide, without eyespots. Peristomium distinctly wider than pro- stomium, composed of 2 apodus segments, their total length about equal to that of setiger 1. Parapodia of anterior 10 setigers reduced, with small post- setal lobes. Parapodia becoming larger, with small presetal lobes becoming evident and postsetal lobes elongate, subconical, slightly longer than setal lobes (Fig. 2e), continuing to posterior end with presetal lobe becoming reduced. Anterior parapodia of adults with 5-6 broadly winged capillary setae; simple bidentate hooded hooks (Fig. 2f) beginning on setiger 23-27 of adults and on setiger 3 of juvenile; setae of middle segments about 5, 2- 3 capillaries, 2-3 hooks; hooks gradually replacing capillaries in posterior segments, usually 3 per parapodium. Pygidium somewhat flattened, bul- bous, with dorsal anus, without anal cirri. Mouthparts relatively massive for size of worm, easily visible through integument. Mandibles (Fig. 2d) brown, about as long as maxillary carriers 424 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 50 jum Larsen BY win OZg Fig. 2. Lumbrinerides jonesi: a, Maxillae, dorsal view; b, M II; c, Maxillae, dorsal view, with M III and M IV somewhat flattened; d, Mandibles, ventral view; e, Parapodium from midregion, anterior view; f, Bidentate hooded hook from same. and M I-III, about twice as long as broad, anteriorly broadly notched and flared, posterior half fused, with minute posterior notch; anterior end with several semicircular lines ventrally, dorsally with lines darkened near an- terior edge and converging posteriorly toward middle; posterior end thick- ened, about twice as thick as wide. Maxillae (Figs. 2a—c) with broad, stout subtriangular carriers together forming obtuse angle open to posterior, more or less constricted posteriorly. M I stout, flared posterolaterally, without accessory teeth; strong, iridescent ligament between their bases. M II (Fig. 2b) broad, stout, with 3 rounded teeth on posterior %4, larger one near middle, smaller ones more posteriad, with posterior elevation; tips of M I dorsally overlying depressed anterior third of each M II. M III with wide, VOLUME 92, NUMBER 3 425 sharp, posterior borders, stout, transversely elongate, about same width as length of M II, probably oriented dorsoventrally when maxillae not everted, with round-tipped, conical tooth. M IV longer than wide, thin, with posterior borders about half as long as those of M III. Maxillary pieces held together by stout ligament-like membrane. Lateral supports extending from middle of M I to M III; M III with small, triangular, ventral supports; M I and M II somewhat calcified. Remarks.—L. jonesi differs from others in the genus principally in struc- ture of Maxillae I and II. The former lack accessory teeth; the latter have 3 molar-like teeth. The specimen referred by Ramos (1976:105) to Lumbri- neris acuta, referred herein to Lumbrinerides sp., is similar to L. jonesi but differs in the dentition of Maxillae II and in shape of the posterior end of the fused mandibles. Etymology.—The species is named in honor of Dr. Meredith L. Jones, who assisted in identification of certain taxa and whose research has con- tributed greatly to knowledge of the Polychaeta. Genus Lumbrineris Blainville, 1828 Lumbrineris dubeni (Kinberg, 1865) Lumbriconereis dubeni Kinberg, 1865:570. Lumbriconereis heteropoda.—Crossland, 1924:4, text-figs. 2-4 [in part, Suez Bay specimens] [not Marenzeller, 1879]. Lumbrineris dubeni.—Hartman, 1948:96. Material examined.—MOZAMBIQUE: Diben, col., 1845; type-speci- men(s), anterior and 3 middle fragments (NRS 382).—SUEZ BAY: 8.2 m; mud; C. Crossland, col., 1904; anterior and 3 middle fragments (BMNH 1923:11:12:7-9). Description.—Colorless in alcohol. Anterior end about 100 segments, middle fragments about 170 segments, together about 250 mm long. Anterior piece 4 mm wide including parapodia. Prostomium longer than wide, blunt- tipped, conical. Peristomium composed of 2 apodus segments, each about as long as setiger 1. Parapodia sub-biramous, with notopodial rudiment at upper border of setal lobe supported by slender internal acicula. Anterior parapodia with short presetal lobe, relatively short, rounded setal lobe hav- ing 4 neuroacicula, notopodial rudiment having 12 notoacicula, and postsetal lobe extending about twice as far as setal lobe. Setal and postsetal lobes lengthening posteriorly; postsetal lobes tapered, becoming more elevated to about 45° angle or bent posteriorly, slightly flattened, with tips extending well past setae by setiger 50; presetal lobes of same short, rounded; setal lobes bluntly conical. All parapodial lobes similar after setigers 50—60 (sim- ilar to those figured by Crossland, 1924:8, text-fig. 6). Anterior parapodia 426 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 4 50 um wit OO! Cc d Fig. 3. Lumbrineris crosslandi (holotype BMNH ZB 1979:1): a, Parapodium of setiger 5, an- terior view; b, Parapodium of setiger 150, anterior view; c, d, Hooded hooks from middle parapodium. with about 12 winged capillaries, gradually reduced to about 5 on setiger 36 and to 1 by about setiger 50, continuing to posterior end of anterior piece (about setiger 100), absent from all segments of middle fragments. Hooded hooks beginning on setiger 36, rapidly replacing all but 1 capillary poste- riorly. Middle parapodia with 3—4 hooks, 2 neuroacicula, about 4 slender notoacicula in notopodial rudiment. Middle fragment of 38 segments appar- ently from near posterior end, with single neuroaciculum in posterior part. Acicula yellow. Mandibles as figured by Crossland (1924:6, text-figs. 3, 4), with broad ligament-like membrane attached to anterolateral edges. Maxillae as figured by Crossland (1924:5, text-fig. 2); M I falcate hooks reaching to anterior ends of M II; M II subsymmetrical, straight, thin pieces with 4 teeth, right with broad space between distal and subdistal tooth, left with distal tooth slightly reduced; M III with 2 pointed teeth, posterior about half length of anterior; M IV with 1 pointed tooth. Maxillary carriers about as long as M II, constricted slightly anterior to middle. Lateral supports between poste- VOLUME 92, NUMBER 3 427 rior third of M I and M III; supports opposite toothed edge of M II subrect- angular, without anterior papilliform projection; brown reinforced area dor- solateral to M III and IV. Remarks.—The specimen(s) from Suez Bay is apparently L. dubeni. Mouthparts, figured by Crossland (1924:5, text-figs. 2-4), are absent, but those of the type specimen(s) are the same as his illustrations of the Suez specimen(s). Winged capillaries occur on all 160 setigers of the anterior fragment, on two of the middle fragments and on most of the third fragment. There are at most 2 hooks on middle and posterior segments of the Suez Bay specimen(s) and up to 4 on the type-specimen(s). The latter are not accompanied by winged capillaries on posterior segments. However, these differences appear to be subspecific. Specimens of L. dubeni may be included in Day’s account of L. heter- opoda heteropoda from southern Africa (Day, 1967:440). Day’s description is not in agreement with that given for L. heteropoda (Marenzeller) by Imajima and Higuchi (1975:30) from Japan. Other possible synonyms of L. dubeni are discussed by Hartman (1948). Lumbrineris crosslandi, new species a Figs. 3, 4 Lumbriconereis heteropoda.—Crossland, 1924:4, figs. 1, 5-7 [in part, Red Sea and Zanzibar specimens] [not Marenzeller, 1879]. Material examined.—RED SEA: Agig Bay, 8.2 m; among Polyzoa; C. Crossland, col., 1904—1905; holotype, complete specimen in 3 pieces (BMNH ZB 1979:1).—Reefs of Shubuk Suakin, 16.5 m, mud (Crossland, 1924:6); C. Crossland, col., 1904-1905; 2 paratypes, 2 anterior and 9 middle fragments (BMNH 1923:11:12:13—20)—ZANZIBAR: Chuaka Bay, inshore, sand; C. Crossland, col., 1901—1902; 2 paratypes, 2 anterior and 2 middle fragments (BMNH 1924:3:1:55-57). Description.—One specimen (BMNH 1923:11:12:13—20) possibly greater than 200 mm long, 6 mm wide including parapodia; holotype apparently complete in 3 pieces, 160 mm long, 4 mm wide, 275 segments. Prostomium somewhat flattened, bluntly conical, slightly longer than wide, without eyes. Peristomium of 2 apodus segments, anterior longer, posterior shorter than setiger 1. Anterior few setigers about 10 times wider than long, lengthening to 5 times wider than long at about setiger 30, continuing to near pygidium with segments again becoming very short. Parapodia sub-biramous with notopodial rudiment on upper side supported by slender acicula. Anterior parapodia including postsetal lobes about '/,) as long as segmental width, setal and postsetal lobes increasing in length to about 14 as long as segmental width at beginning of middle third of body, continuing about as long to 428 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 4. Lumbrineris crosslandi (holotype BMNH ZB 1979:1): a, Maxillae, dorsal view; b, Right maxillae II-IV, ventral view; c, Mandibles, dorsal view; d, Same, ventral view. posterior end. Setiger 5 parapodium (Fig. 3a) with 6 notoacicula; setal lobe short, about as long as wide, supported by 2 relatively slender yellow aci- cula; presetal lobe rounded, relatively well developed; postsetal lobe slightly longer, more slender than setal lobe. Setiger 150 parapodium (Fig. 3b) with VOLUME 92, NUMBER 3 429 notopodial rudiment supported by 4 slender notoacicula; setal lobe subcon- ical, supported by 2 stout acicula; presetal lobe well developed; postsetal lobe elongate, elevated at about 45° angle or bent posteriorly. Only winged capillaries in about anterior 40 setigers (37-45); transitional region contain- ing both winged capillaries and simple hooded hooks extending to about setiger 90 (54-93, setiger 54 on holotype, but almost all setae broken off in midregion, thus possibly extending farther posteriorly); posterior 7% of body with hooks only (Figs. 3c, d). Anterior parapodia with 7-8 limbate setae; about 4 hooks on parapodia of posterior 74. Pygdium cylindrical, slightly longer than wide, with terminal anus surrounded by 4 long, conical anal cirri. Mandibles (Figs. 4c, d) broad anteriorly, gradually reduced to less than 14, total width on posterior third, narrowly divided anteriorly and posteriorly, with ventral anterior edge broadly rounded, without attaching ligaments. Maxillae symmetrical (Figs. 4a, b). M I long, falcate, not reaching to anterior borders of M II; M II strongly curved around anterior ends of M I, with 5 teeth, anterior one very small, and anterior rounded boss opposite toothed edge; M III with 2 teeth, anterior tooth longer than posterior but both rel- atively well developed (tips apparently broken off); M IV with single tooth. Tips of teeth of M II-IV clear in glycerin, probably calcified in life. Lateral support from M I to base of M III; rounded support between M I and M II; support opposite edge of M II subrectangular, without anterior papilliform projection. Remarks.—Crossland’s text-figures 1—7 suggest that his specimens com- prised 2 species. Mouthparts of his identified specimens in the British Mu- seum are now missing. However, included in a lot of 2 Lumbrineris speci- mens from among Crossland’s unworked material from the Red Sea was a complete specimen that agreed with his other specimens from the Red Sea and Zanzibar and is herein designated the holotype. The other species in Crossland’s identified material is L. dubeni (Kinberg). Specimens of L. crosslandi may also be included with specimens of L. dubeni in Day’s (1967) account of L. heteropoda heteropoda from southern Africa. Etymology.—The species is named in honor of Dr. Cyril Crossland, who collected and reported the specimens from the Red Sea and tropical East Africa. Lumbrineris ernesti, new species Figs. 5, 6 Lumbrinereis heteropoda.—Monro, 1933a:258, fig. 8 [not Lumbriconereis heteropoda Marenzeller, 1879]. Lumbrineris erecta.—TYaylor, 1971:354.—Hall and Saloman, 1975:14 [list] [not Lumbriconereis erecta Moore, 1904]. Material examined.—FLORIDA: Anclote Anchorage, Tarpon Springs, 28°12'39"N, 82°47'39"W, 3.5 m, mixed seagrasses, fine sand; R. Ernest and 430 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 10 pm MM Fig. 5. Lumbrineris ernesti: a, Parapodium of setiger 20, anterior view; b, Parapodium from near pygidium, anterior view; c, Largest hooded hook of middle parapodium; d, Middle hook from same; e, Hook with broken tip; f, Hook from setiger 35 showing hoods and curvature. J. Studt, cols., 5 Dec. 1975; holotype (USNM 57438); 2 paratypes (AHF POLY 1269; FSBC I 20451).—Hutchinson Island, Sta. I, 27°21.3'N, 80°14.1'W, about 8 m, fine quartose sand; 2 paratypes (FSBC I 20452, 20453).—Sta. IV, 27°20.7'N, 80°12.8'W, about 11 m, coarse calcareous VOLUME 92, NUMBER 3 431 sand; 2 paratypes (FSBC I 20454, 20455).—Dry Tortugas Ecological Survey Station 153; Coleman and Tandy, cols.; 1 paratype (BMNH 1932:12:22:44— 45).—Lower Tampa Bay, 27°35'26"N, 82°45'26’W, 3 m; J. Taylor and C. Saloman, cols., 1963; 2 paratypes (ZMH P-15538; FSBC I 13183).—Tampa Bay, J. Taylor, col., 1963; 7 paratypes (USNM 45598).—MISSISSIPPI: 29°55'N, 88°43.5’'W, 15 m, fine sand, silt, BLM Sta. 1 F; R/V Colombus Iselin, May 1974; 1 paratype (USNM 55878).—NORTH CAROLINA: off Beaufort, 34°29'N, 76°13’W, about 33 m; F. Grassle, col.; 1 paratype (USNM 54280). Description.—Colorless in alcohol. Length up to 105 mm, width 5 mm including parapodia, 270 segments. Prostomium somewhat flattened, sub- conical, slightly longer than wide. Peristomium with 2 apodus segments. Body tapered anteriorly for about first 20 segments; segments about 8 times wider than long at setiger 20; parapodia sub-biramous with notopodial ru- diment; setal lobe cylindrical; postsetal lobes about as long as setal lobes in anterior segments (Fig. 5a); 3 yellow acicula; notopodial rudiment with 5—6 slender notoacicula. Parapodial lobes becoming more elongate in middle and posterior segments (Fig. 5b). Capillary setae only for first 42-47 setigers of larger specimens, first 17-18 setigers of small specimens; maximum of 11 per parapodium in anterior region; capillary setae continuing to posterior end, except on 2 juveniles lacking such after setiger 40. Hooded hooks beginning on setiger 18—19 of juveniles, on setiger 47 of largest specimens, continuing to posterior end; parapodia of posterior segments with 3—4 hooks, 1 capillary, 2-3 notoacicula, single neuroaciculum. Hooks with large main tooth and 12-16 secondary teeth decreasing in width and increasing in length toward apex (Figs. Sc, d); tips of hooks of large specimens often damaged or worn; shafts only slightly bent (Figs. 5b, f). Pygidium cylindri- cal, short, with terminal anus surrounded by 4 conical anal cirri about twice as long as postsetal lobes. Mandibles flared anteriorly, medially concave; broad ligament-like mem- branes attached to anterolateral edges; mandibles broadly divided for short distance posteriorly. Maxillae (Figs. 6a—c) symmetrical with carriers almost as long as M I, about twice as long as broad, strongly constricted near anterior end, acutely pointed posteriorly; anterior margins nearly perpen- dicular to long axis. M I strongly falcate, round-tipped, reaching to anterior end of M II; M II (4 + 4 — 5), mght with broad space between distal and subdistal tooth of adults, broad space replaced by small tooth in small spec- imens, large gravid specimen (USNM 55878) with additional incipient small tooth basally, left with distal tooth often reduced; calcified areas often on anterior edges of teeth (clear in glycerin), pieces almost straight, not partic- ularly curved around anterior borders of M I; M III with 2 teeth, distal one pointed, proximal one rounded, often reduced to right-angular corner; M IV with single, acutely pointed tooth and small calcified areas slightly below 432 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 6. Lumbrineris ernesti: a, Maxillae, flattened, dorsal view; b, Right maxillae II-IV, ventral view; c, Maxillae, slightly flattened, dorsal view; d, Mandibles, dorsal view; e, Same, ventral view; f, Posterior end, dorsal view; g, Anterior end. tip (clear in glycerin). Maxillae with several auxillary supports, lateral one apparently articulated near anterior end, extending from about posterior third of M I to M III; support opposite toothed edge of M II semi-oval, extending almost full length with anterior papilliform extension; support VOLUME 92, NUMBER 3 433 between posterior half of M I and posterior third of M II flattened, anteriorly rounded; brown, suboval area dorsolateral to M III and M IV;; calcified area often visible posterior to M II. Remarks.—The specimen from Tortugas, Florida (BMNH 1932:12:22:44~— 45), reported by Monro (1933a) as L. heteropoda, agrees with L. ernesti. Monro (1933b) reported L. heteropoda from the Galapagos and Pacific Pan- ama (Balboa); they appear to be a different species; the Balboa specimen was referred to L. erecta by Fauchald (1977). L. heteropoda (Marenzeller, 1879) has almost completely divided mandibles, symmetrical maxillae with M III having 2 subequal teeth, and parapodia with short postsetal lobes in the midregion and short, elevated postsetal lobes in the posterior region (Imajima and Higuchi, 1975). Specimens from Tampa Bay, Florida, reported by Taylor (1971) and Hall and Saloman (1975) as Lumbrineris erecta (Moore, 1904), also agree with L. ernesti. L. erecta is a similar species, but differs from L. ernesti in having erect postsetal lobes in posterior segments, and in the shape and dentition of the hooded hooks (see Moore, 1904, and Fauchald, 1970). Lumbrineris ernesti as defined herein is similar to L. dubeni, differing primarily in the shapes of M II and III. M III of L. dubeni have 2 pointed teeth, and those of L. ernesti have 1 pointed tooth and 1 rounded tooth which is often only a right-angular corner. M II of L. ernesti have a rounded papilliform extension on supports opposite the toothed edge which is absent on L. dubeni. Hooded hooks of both species are very similar. Etymology.—The species is named for Mr. Robert G. Ernest, who col- lected some of the specimens. Lumbrineris tenuis (Verrill, 1873) Fig. 7 Lumbriconereis tenuis Verrill, 1873:594. Lumbrineris tenuis. —Hartman, 1942b:54; 1944b:340, pl. 49, figs. 3-5.—Pet- tibone, 1963:264 [in part], figs. 70a—e, ?f—Day, 1973:62.—Gardiner, 1976:199, figs. 26p-r. Lumbrineris bassi Hartman, 1944a:150, pl. 10, figs. 217-223; 1951:58.—Tay- lor, 1971:347.—Hall and Saloman, 1975:14 [list]. Material examined. —CONNECTICUT: Stony Creek, E of New Haven; A. E. Verrill, col., May 1872; syntype, anterior end and 2 middle fragments (YPM 2733)—MASSACHUSETTS: Vineyard Sound; U.S. Fish Comm., 1883; 12 specimens (USNM 13121)—NEW YORK: Long Island Sound; U.S. Fish Comm., 1874; 2 specimens (USNM 13123).—NORTH CARO- LINA: off Beaufort, 34°34’N, 76°25’W, 19 m, sand and shell; J. H. Day, col., 4 June 1965; 84 specimens (USNM 55173).—White Oak River, Swans- boro, intertidal, muddy sand; H. Lee, col., Aug. 1972; 1 specimen (USNM 434 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 25 ym Fig. 7. Lumbrineris tenuis: a, Posterior parapodium, anterior view (USNM 13121); b, Same (FSBC I 18972); c-e, Hooded hooks; c, Setiger 14 (USNM 13121); d, Same, posterior para- podium; e, Posterior parapodium (FSBC I 18972); f, Maxillae (USNM 51465); g, Left M II of same, lateral view; h, Maxillae (FSBC I 18972); i, Right M II-M IV of another specimen, medial view (FSBC I 18972); j, Left M II of same, lateral view. 53016).— FLORIDA: Biscayne Bay, Miami; R. Rosenberg, col., 1974; 1 specimen (USNM 51465).—Tampa Bay, 27°45'40"N, 82°44'26"W, | m, sand, algae and seagrass cover; J. Taylor and C. Saloman, cols., 23 Sept. 1963; | specimen (FSBC I 13330).—27°43'26’N, 82°43'47”W, 1 m, sand, algae and VOLUME 92, NUMBER 3 435 seagrass cover; J. Taylor and C. Saloman, cols., 1 Oct. 1963; 26 specimens (USNM 57439; FSBC I 18972).—27°41'34"N, 82°44'03”W, 1 m, sand, algae and seagrass cover; J. Taylor and C. Saloman, cols., 15 Oct. 1963; 1 spec- imen (FSBC I 18929).—Anclote Anchorage, Tarpon Springs, 28°12.5'N, 82°46.5'W, 0.5 m, sand, J. Studt and R. Ernest, cols., 5 Dec. 1975; 4 spec- imens (FSBC I 17384). Description.—Generally uncolored, occasionally with scattered brown pigment on prostomium and anterior segments. Maximum length 350 mm (in life), 1 mm wide, 200 segments (Verrill, 1873; Pettibone, 1963). Prosto- mium slightly flattened, bluntly conical, about as long as wide. Peristomium of 2 apodus segments, each about as long as first setiger. Parapodia unira- mous, with 3 slender yellow acicula in setal lobes of anterior parapodia, single stouter one in posterior parapodia. Setal lobes short on anterior few segments, longer on setigers 8—15, bluntly conical, gradually shorter toward middle, gradually lengthening on posterior half and becoming subcylindrical. Presetal lobes short, rounded throughout. Postsetal lobes of anterior seg- ments extending about twice as far as setal lobes, subtriangular to auricular, becoming slightly elevated, shorter, more slender toward middle, gradually lengthening, becoming digitiform to club-shaped near pygidium (Figs. 7a, b), often elevated dorsally or bent anteriorly or posteriorly. About 10-17 anterior setigers with up to 10 limbate capillary setae; simple hooded hooks (Figs. 7c, e) gradually replacing capillaries by about setiger 30; 2-3 hooks on middle and posterior parapodia. Capillary setae longer, broader on an- terior segments, becoming shorter, more slender after appearance of hooks. First hooded hooks with longer shafts, fewer apical teeth, otherwise similar to more posterior hooks. Tips of hooks of anterior segments with pointed primary tooth, 5—6 secondary teeth decreasing in length and thickness api- cally; hooks of middle and posterior segments slightly stouter, more strongly curved, often with irregular primary tooth and 10-12 secondary teeth de- creasing in length and thickness. Pygidium with 2 pairs of digitiform to conical anal cirri about as long as width of pygidium. Mandibles thin, almost translucent in glycerin, anteriorly flared, poste- riorly divided up to about half total length as figured by Verrill in Hartman (1944b, pl. 49, fig. 5b). Maxillae as figured (Figs. 7f-j). Carriers brown, shorter than M I, subtriangular, slightly concave laterally. M I light brown, broad, thin, with denticulate border below short, falcate tip; M II light brown, not particularly stout, with 4—6 teeth, left often with one more tooth than right; M III brown, with single, dark, upper tooth extending as broad, dark, posterior border below; M IV with dark, conical to somewhat rounded tooth. Light brown lateral support from M I to anterior base of M II and base of M III; area of dark brown cells between bases of M I and M II; short, suboval, dark brown support on medial border of M II opposite teeth; 436 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON dark support below medial border of M III; larger, dark support on antero- medial border of M IV connected by narrow isthmus to light brown dor- solateral support. Remarks.—The syntype material contains an anterior end of about 25 segments of L. tenuis and 2 middle pieces which may be the same species. However, the greater part of the material consists of middle pieces of a Drilonereis. The latter probably accounts for Hartman’s statement (1942b:54) that ‘“‘parapodial lobes are short throughout, inconspicuous.”’ Verrill (1873) described slender hooded hooks beginning on setiger 9 [probably a large specimen?], changing to stouter, recurved hooks with 2 or 3 hook-like denticles on the end beginning at setiger 17. The latter may possibly be the same as figured by Pettibone (1963:264, fig. 70f), but hooded hooks appear at setiger 15 on the syntype I examined (YPM 2733) and are the same as I have figured from other specimens (Figs. 7c—e). Posterior segments are absent on the syntype, but those figured by Verrill in Hartman - (1944b:pl. 49, fig. 3a”) are not dissimilar from parapodia of most specimens I have examined, and are in fact quite similar to those of specimens from off Beaufort, North Carolina (USNM 55173). Posterior parapodia of the latter are slightly shorter than those figured (Figs. 7a, b). Maxillae II of a large specimen from off Beaufort have 6 teeth on the left and 5 on the right. This is also the case on the syntype, and is the opposite of the number of teeth on left and right pieces given by Hartman (1942b). No hooded hooks were found before setiger 11 on any specimen, including many very small ones, and it is my opinion that they do not occur before setiger 9 or 10 in the species. Further, anterior hooded hooks do not differ greatly in shape from those farther back. This is in disagreement with the description and figures of Pettibone (1963:264, figs. 70h-j), probably result- ing from her inclusion of L. hebes as a synonym of L. tenuis. Day (1973) stated that mandibular shafts were in contact throughout, but I cannot confirm his observation. Lumbrineris hebes (Verrill, 1879) Figs. 8, 9a—c Lumbriconereis obtusa Verrill, 1874:383 [not Kinberg, 1865]. Lumbrinereis hebes Verrill, 1879:174. Lumbrineris hebes.—Hartman, 1942b:9. Lumbrineris tenuis.—Pettibone, 1963:264 [in part, not Lumbrinconereis ten- uis Verrill, 1873], figs. 70g-}. Material examined.—MAINE: Casco Bay, 13 m; U.S. Fish Comm., 7 Aug. 1873; 6 syntypes (USNM 15814).—MASSACHUSETTS: Quincy Bay, mid-way between West Head and Sunken Ledge, 5-6 m; J. Clark and class, cols., 8 May 1967; 1 specimen (USNM 50542). VOLUME 92, NUMBER 3 437 De) e) Oo = 3 win GZ Cc ,__200 jim Fig. 8. Lumbrineris hebes: a-c, Parapodia, anterior view (USNM 50542): a, Setiger 5; b, Setiger 125; c, Setiger 22; d, Pygidium, ventrolateral view, syntype (USNM 15814); e-g, Hood- ed hooks (USNM 50542): e, Setiger 6; f, Setiger 22; g, Posterior setiger. Description.—Syntypes colorless; other specimen with numerous irreg- ularly arranged pigment spots dorsally and ventrally on prostomium and anterior segments. Length about 25 mm, width about 0.75 mm (Verrill, 1874); largest specimen (USNM 50542) incomplete posteriorly, 32 mm long, 1.3 mm wide including parapodia, with about 130 segments. Prostomium about as long as wide, slightly flattened, bluntly conical. Peristomium of 2 apodus segments, each slightly shorter than setiger 1. Parapodia uniramous 438 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON (Figs. 8a—c), with 3 slender yellow acicula in setal lobes of anterior seg- ments, single stouter aciculum in posterior segments. Setal lobes short, bluntly conical on anterior segments, about twice as long on posterior seg- ments, subconical. Presetal lobes short, rounded on anterior parapodia, be- coming pointed on posterior parapodia. Postsetal lobes of anterior segments broad, with rounded tips extending about twice as far as setal lobes; lobes on middle segments shorter, elevated dorsally; lobes on posterior segments long, flattened, elevated at greater than 45° angle or bent backward. Anterior parapodia with 2—4 upper limbate capillaries, 2—4 slender, simple, hooded hooks with long hoods (Fig. 8e), and short, slender lower limbate capillary; hooks gradually becoming stouter, with shorter hoods after about setiger 10 (Fig. 8f), replacing limbate capillaries by about setiger 25; 2—4 subequal hooks on middle and posterior parapodia, stouter on middle than on pos- terior parapodia (Fig. 8g). Pygidium (Fig. 8d) with 4 rounded, posterior lobes, dorsolateral pair shorter, broader, ventral pair longer; anus opening between dorsolateral pair; no anal cirri. Mandibles (Fig. 9b) thin, translucent in glycerin, anteriorly flared, pos- teriorly divided for about % total length. Maxillae as figured (Figs. 9a, c). Carriers shorter than M I, thin, brown; M I brown, with smooth border below falcate tip; M II brown, subequal, straight, stout, with 4 teeth, right one with stout distal tooth; M III dark brown, subtriangular, with rounded tooth above dark posterior border; M IV dark conical tooth. Dark brown lateral support between M I and anterior base of M II and base of M III; area of dark cells between posterior parts of M I and M II; long, dark, subrectangular support with anterior extension on medial border of M II opposite teeth; dark support medially below M III; dark brown, subtrian- gular support anterior to M IV; light brown support dorsolateral to M IV, separated from tooth and anterior support. Remarks.—Lumbrineris hebes was originally described as Lumbriconer- eis obtusa, a homonym preoccupied by Kinberg, 1865; the new name was proposed by Verrill in 1879. Lumbrineris hebes was referred to L. tenuis by Pettibone (1963) but it differs from L. tenuis in having: 1) hooded hooks which begin on the first setiger and which differ in shape from those of middle and posterior setigers; 2) a pygidium with four rounded lobes and no anal cirri; 3) M I with a smooth border below the falcate tip; 4) prominent supports lateral to M I; 5) longer and darker supports opposite the toothed border of M II; and 6) M IV with a large anterior support separated from the dorsolateral support. Specimens of L. hebes do not appear to be juve- niles of L. tenuis. Many smaller specimens of L. tenuis were examined; none had hooks appearing before setiger 10, while all specimens of L. hebes had hooks appearing on setiger 1. The seta described by Verrill (1874) as ‘‘long and slender, with a very slender setiform tip’? may be the lower capillary seta of anterior parapodia. VOLUME 92, NUMBER 3 439 Lumbrineris hebes was also reported from Noank Harbor, Connecticut by Verrill (1875), Eastport, Maine by Webster and Benedict (1887), and Labrador by Moore (1909). However, specimens originally identified by Verrill as Lumbriconereis obtusa and Lumbrinereis hebes from south of Cape Cod (USNM 13121, 13123) have proven to be L. tenuis, and remains of Webster and Benedict’s and Moore’s specimens deposited in the Smith- sonian Institution are middle pieces (Dr. Pettibone, personal communica- tion) insufficient to confirm their records. Moore (1911:290) indicated that the “‘jaws’’ of L. hebes were similar to those of L. inflata (Moore, 1911). However, M III of L. inflata are multi- dentate and M IV are bidentate, demonstrating that L. inflata is certainly a different species. Fauchald (1970:215) apparently followed Moore’s de- scription of the maxillae but also incorrectly attributed the composite hooks of L. inflata to L. hebes. A species which may be the same as that reported by Moore (1909) was reported from 40-200 m off Beaufort, North Carolina, by Day (1973:60) as Lumbrineris sp. (USNM 51147). The latter has tri- and quadri-dentate M III and uni- and bidentate M IV. Lumbrineris testudinum (Augener, 1922) Figs. 9d—h Lumbriconereis testudinum Augener, 1922:46.—Hartman, 1959:331. Lumbrineris testudinum.—Hartman, 1959:337. Material examined.—FLORIDA: Tortugas, Southwest Channel; Hart- meyer, col.; holotype, possibly sexually mature male (ZMB 6399). Description.—Colorless in alcohol. About 14 mm long, slender, 93 setig- erous segments. Prostomium about as wide as long, almost globular. Per- istomium of 2 apodus segments, each about as long as setiger 1. Parapodia uniramous, with about 3 yellow acicula in setal lobes of anterior segments, single stouter aciculum in posterior segments. Setal lobes of anterior seg- ments (Fig. 9d) bluntly conical, short on first few segments, slightly longer on setiger 10, gradually reduced in length and about as wide as long pos- teriorly. Presetal lobes short, rounded throughout. Postsetal lobes subtrian- gular, extending almost twice as far as setal lobes on anterior segments; short, rounded, only slightly longer than presetal lobes on middle and pos- terior parapodia. Simple hooded hooks and limbate capillary setae beginning with first setiger; about 3 hooks on middle and posterior parapodia. Hooks of anterior segments (Fig. 9f) stout, with 4—S subequal teeth; hooks of middle and posterior segments (Fig. 9g) similar to anterior hooks in general shape, with primary tooth surmounted by 8—10 secondary teeth decreasing in length and stoutness apically. Pygidium with pair of long, conical dorsal cirri and pair of short, conical ventral cirri surrounding anus. 440 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON wit 002 wit O02 win G2 we g Fo e 25 ym Fig. 9. Lumbrineris hebes (USNM 50542): a, Maxillae; b, Mandibles, ventral view; c, Right M II-M IV, medial view. Lumbrineris testudinum, holotype (ZMB 6399): d, Anterior para- podium, anterior view; e, Posterior parapodium, anterior view (not scaled); f, Hooded hook, setiger 1; g, Hooded hook, posterior parapodium; h, Maxillae. Mandibles thin, translucent, anteriorly flared, with dark lateral corners; posterior ends not examined. Maxillae as figured (Fig. 9h). Carriers brown, constricted laterally, shorter than M I. M I very light brown, long, thin, falcate, with smooth border; M II very light brown, thin, laterally curved anteriorly around tip of M I, with 5 teeth; M III light brown, with distinct pointed tooth above broad, posterior, dark brown border; M IV with round- VOLUME 92, NUMBER 3 441 ed brown tooth above darker posterior border. Lateral support between M I and anterior base of M II and base of M III large, similar to those described for L. hebes, formed of distinct, dark brown cells on posterior part; area of dark brown cells between bases of M I and M II; dark brown support on medial side opposite toothed border of M IJ; support medial to tooth of M III; single brown support anterior to tooth of M IV, with dorsolateral sup- port, if present, very indistinct. Remarks .—Stout, simple hooded hooks beginning on the first setiger dis- tinguish the species from other similar small, slender species with uniramous parapodia and short parapodial lobes reported herein. Additionally, M III has a single pointed tooth; L. parvapedata (Treadwell, 1901), a much larger species, has similar parapodia and setae, but has 2 teeth on M III. Lumbrineris verrilli, new species Figs. 10, 11 Lumbrineris sp. Taylor, 1971:363, figs. 7a, b.—Hall and Saloman, 1975:14 [list]. Material examinéd.—FLORIDA: Lower Tampa Bay (National Marine Fisheries Service Tampa Bay Study; J. Taylor and C. Saloman, cols.), 27°33'27'N, 82°42'36"W, 5 m, sand; 4 Nov. 1963; holotype (USNM 57440), 28 paratypes (USNM 57441).—27°39'19"N, 82°42'10"W, 4 m, sand; 15 Oct. 1963; 2 paratypes (FSBC I 20459).—27°37'5S’N, 82°41'30"W, 4 m, sand; 18 Oct. 1963; 1 paratype (FSBC I 20460).—27°34'34’N, 82°39'58”W, 4 m, sand; 28 Oct. 1963; 5 paratypes (FSBC I 20461).—27°34’04"N, 82°39'42”W, 5 m, sand; 28 Oct. 1963; 13 paratypes (FSBC I 20462).—27°33'03’N, 82°39'12’W, 4 m, sand; 6 Nov. 1963; 29 paratypes (FSBC I 20463).—27°34'54’N, 82°43’01"W, 7 m, sand; 30 Oct. 1963; 24 paratypes (AHF POLY 1270).— 27°34'24"N, 82°42'53"W, 6 m, sand; 4 Nov. 1963; 4 paratypes (SIFP 50:0781).—27°33'S57"N, 82°42'46"W, 5 m, sand; 4 Nov. 1963; 89 paratypes (FSBC I 20464).—27°32'53”"N, 82°42'27"W, 3 m, sand; 4 Nov. 1963; 21 para- types (ZMH P-15539).—27°31'41"N, 82°42'10"W, 3 m, sand; 19 Nov. 1963; 1 paratype (FSBC I 20465).—27°41'46’N, 82°43'34"W, 7 m, sand; 18 Nov. 1963; 1 paratype (FSBC I 20466).—Tampa Bay; J. Taylor, col., 1963; 10 paratypes (USNM 45701).—Hutchinson Island, Sta. IV, 27°20.7'N, 80°12.8'W, about 11 m, coarse calcareous sand; | paratype (FSBC I 18860).—Sta. V, 27°22.9'N, 80°13.9’W, about 11 m, coarse calcareous sand; 11 paratypes (USNM 57443; FSBC I 18861, 18863-6).—Biscayne Bay, Miami; R. Rosenberg, col., 1974; 39 paratypes (USNM 57442). Description.—Many specimens in thin, mucoid tubes covered with detri- tis. Body white to light reddish-orange in alcohol. Largest specimen (USNM 57442) 45 mm long, 0.7 mm wide including parapodia, about 110 segments, incomplete; holotype 20 mm long, 0.4 mm wide including parapodia, 160 442 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Ss wit OZ wit OZ (Sees wrt OZ Fig. 10. Lumbrineris verrilli, a-f, Hutchinson Island specimens: a, Anterior parapodium, anterior view; b, Parapodium of midregion, anterior view; c, Posterior parapodium, anterior view; d, Hooded hook, anterior parapodium; e, Lower hooded hook, middle parapodium; f, Upper hooded hook, middle parapodium; g, Biscayne Bay specimen (USNM 57442), mandi- bles, ventral view. segments, with regenerating pygidium. Prostomium acorn-shaped to bluntly conical, about as long as wide, tip often slightly acuminate. Peristomium of 2 well-marked apodus segments, each about as long as setiger 1. First few segments slightly narrower and shorter than those following; segments wid- est at about setiger 10, gradually decreasing in width posteriorly; middle segments half to about as long as wide; segments gradually shortening pos- teriorly. Parapodia uniramous, well separated, short, conical, truncate, about as thick as long on anterior and middle segments, slightly longer posteriorly; 3 yellow acicula in parapodia of anterior and middle segments, single aciculum in posterior parapodia. First few pairs of parapodia slightly smaller than following, with short, rounded postsetal lobes; presetal lobes VOLUME 92, NUMBER 3 443 200 ym Fig. 11. Lumbrineris verrilli, a-c, Hutchinson Island specimens: a, Anterior end, dorsal view; b, Posterior ends, dorsal view; c, Maxillae from whole mount; d-f, Tampa Bay speci- mens: d, Mandibles, ventral view; e, Maxillae of small specimen; f, Right M III, M IV of e, flattened; g-i, Biscayne Bay specimen (USNM 57442): g, Maxillae of large specimen; h, Left M II-M IV, medial view; i, Right M III, M IV, flattened. absent or reduced. Anterior parapodia largest on about setiger 10, with long, broad postsetal lobes, gradually reduced posteriorly, with short presetal lobes developing on about setiger 15. Parapodia of midregion shorter than anterior parapodia, with short, rounded presetal lobes, slightly longer, dig- itiform, somewhat elevated postsetal lobes. Parapodia gradually lengthening in posterior region, with reduced presetal lobes and narrow, cylindrical, slightly elevated postsetal lobes extending to about apices of hooks. Ante- rior segments with 3-4 upper winged capillary setae, 1-2 simple hooded hooks beginning on setiger 1, and single lower capillary; winged capillaries often shorter on anterior 4 setigers, reduced in number after setiger 10, 444 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON absent after about setiger 25; hooded hooks of anterior segments with much longer hoods, about half thickness of following hooks, slightly geniculate; apices of hooks forming acute angle with about 5 subequal secondary teeth above slightly larger primary tooth. Middle and posterior segments with about 3 hooded hooks, uppermost stouter than others; hooks with shorter hoods, stouter tips, large, pointed primary tooth and about 8 secondary teeth gradually decreasing in length and thickness. Pygidium with terminal anus; single pair of divergent, tapering cirri about as long as width of py- gidium. Mandibles (Figs. 10g, 11d) fragile, light brown with dark anterior borders and clear posterior sides in larger specimens, almost completely translucent in glycerin in smaller specimens, anteriorly flared, posteriorly fused, at least in larger specimens. Maxillae as figured (Figs. llc, e-i). Carriers dark brown, formed in part of distinct hexagonal cells in larger specimens, to- gether slightly longer than broad, about as long as M J, slightly rounded anteriorly, slightly constricted laterally, pointed posteriorly. M I light brown, often with darker posterior corners and tip, relatively stout in larger specimens (Fig. 11g), with broad, thin interior border in smaller specimens (Figs. llc, e); interior border occasionally denticulate, possibly from dam- age or wear, with short, falcate tip; M II light brown, very stout, with 3-4 dark, stout teeth; M III brown, with single, rounded tooth extending ven- trally as dark, broad posterior border; M IV with single, dark brown, straight, broad posterior tooth. Lateral support from M I to M III extremely reduced; area of dark brown cells between posterior bases of M I and M II; small, brown, semi-oval support on medial side opposite teeth of M II and M III; single large, semi-oval support anterior to tooth of M IV, not divided into dorsolateral and anteromedial areas, with uniformly brown area prox- imal to tooth and cellular area distally in larger specimens, almost uniformly brown in smaller specimens. Remarks.—Lumbrineris verrilli resembles L. tenuis, L. hebes and L. tes- tudinum in being small and slender and in having a similar prostomium and parapodia. It differs from L. tenuis by having slender hooded hooks on the first setiger, anterior hooded hooks different from those of middle and pos- terior setigers, a single pair of anal cirri, a broad, straight tooth with a single support on M IV, and very reduced lateral supports between M I and the base of M III. L. hebes has prominent lateral supports from M I to M III, a pointed tooth on M IV, and no anal cirri. Stout hooded hooks beginning on the first setiger of L. testudinum are similar to those of middle and posterior setigers; M II are thin, with 5 teeth; lateral supports are prominent; and there are 2 pairs of anal cirri. Etymology.—tThe species is named in honor of Prof. A. E. Verrill, who described many polychaete species during his extensive career. VOLUME 92, NUMBER 3 445 Family Arabellidae Hartman, 1944 Genus Arabella Grube, 1850 Subgenus Cenothrix Chamberlin, 1919 KEY TO SPECIES OF ARABELLA (CENOTHRIX) OF THE WESTERN ATLANTIC 1) Maxillae symmetncal, both M Il long... .......5... A. multidentata — Maxillae asymmetrical, left M II short, right M II] long ........... 2 Peelecion lewith bind tip a. hae ee ees oes Aa oe ace cece A. mutans SCE Wiel wath Simple tip) Sen. Peek. oo Pat a. eld A. maculosa Arabella (Cenothrix) mutans (Chamberlin, 1919) Fig. 12 Cenothrix mutans Chamberlin, 1919:330, pl. 61, figs. 2-9, pl. 62, fig. 1. Arabella mutans.—Crossland, 1924:71. Arabella (Cenothrix) asymmetrica.—Orensanz, 1974:387, fig. 2 [not Ara- bella novecrinita var. asymmetrica Crossland, 1924]. Material examined.—EASTER ISLAND: shore; holotype (USNM 19740).—FLORIDA: Hutchinson Island, Sta. II, 27°21.6’N, 80°13.2’W, about 11 m, coarse calcareous sand; 2 specimens (USNM 54694, 54695).— Sta. IV, 27°20.7'N, 80°12.8’W, about 11 m, coarse calcareous sand; 2 spec- imens (USNM 54696; FSBC I 18867).—Sta. V, 27°22.9'N, 80°13.9'W, about 11 m, coarse calcareous sand; 1 specimen (FSBC I 18868).—Florida Middle Ground, 28°35.0’N, 84°14.9’'W, 31 m, on coral; 1 specimen (FSBC I 18869).—_NORTH CAROLINA: off Beaufort, 130 m, mud and sand; J. H. Day, col. and det., 1965; 8 specimens (USNM 51155). Description.—Anterior and posterior ends yellow, other segments with yellowish-orange to dark reddish-brown transverse bands near middle of each segment, areas near segmental constrictions usually colorless or yel- low; large specimen from Florida Middle Ground uniformly very light brown or colorless. Largest Hutchinson Island specimen posteriorly incomplete, longer than 110 mm, 480 segments. Prostomium bluntly conical, with 4 dorsal, posterior eyes. Peristomium of 2 apodus segments shorter than fol- lowing segments. Parapodia similar throughout body, sub-biramous; setal lobes usually with 3 stout, sharply pointed acicula; tips broken off or very thin, projecting through integument; notopodial rudiment of 1-2 small, rounded lobes on dorsal side of setal lobe, with 2-3 slender internal acicula. Postsetal lobe projecting from basal half of setal lobe, dorsoventrally about half as wide and about twice as long as setal lobe, often slightly curved dorsally. Setae of anterior segments winged capillaries. Setae of middle and posterior segments 4—5 of 4 types: 1) upper seta winged capillary with wings smooth or lightly serrate (Fig. 12g); 2) transversely serrate seta below upper 446 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 12. Arabella mutans: a, Maxillae, somewhat schematic; b, Maxillae, flattened; c, Max- illary supports; d, Mandibles; e, Posterior end, lateral view; f, Parapodium of middle segment, anterior view; g, Upper winged capillary seta from middle segment; h, Transversely serrated seta from same; i, Asymmetrically hooded acicular seta from same, posterior view; j, ASym- metrically hooded acicular seta from Hutchinson Island specimen, anterior view; k, Same from holotype, anterior view. (a, d, g, h, j, and k not scaled). VOLUME 92, NUMBER 3 447 winged capillary with several transverse rows of prominent serrations (Fig. 12h); 3) winged capillary seta below transversely serrated setae, shorter, with wings more strongly serrated than upper winged capillary; 4) lower seta stouter than others, acicular, with tip projecting from asymmetrical hood and with hoods visible on both sides of seta (Figs. 12i-k). Pygidium truncate conical, with 4 subequal anal cirri about as long as posterior width of pygidium (Fig. 12c). Maxillae I and II asymmetrical (Figs. 12a, b): MI (2+9-8)+(1+5 — 6); MII (7 + 11); MIII-V symmetrical: M III (1 + 4 — 5); MIV (1 + 4); M V 1. Maxillary carriers about twice as long as maxillae; median unpaired rod extending posteriorly for slightly more than half length of paired car- riers, anterior end truncated, apparently joining ligaments or membranes attached to bases of M II. Mandibles (Fig. 12d) about as long as maxillae, dark, attached by short ligament anterior to mid-transverse line. Remarks.—The holotype of Cenothrix mutans appears indistinguishable from Hutchinson Island specimens in most respects. Color is not as prom- inent on the former, shape and length of the median maxillary carrier could not be determined without damaging the specimen, and mandibles are about 1.3 times longer than maxillae. Chamberlin stated that eyes were absent on the holotype; they are present but covered by the first peristomial segment. Maxillae I were incorrectly figured by Chamberlin; the left M I has a bifid falcate tip, and the right M I has a simple falcate tip. At least the 4 larger specimens from North Carolina (USNM 51155) are the same species. The species may also have been partly described and figured by Crossland (1924:82, specimen no. W.9.2 only) as a variant specimen of Arabella no- vecrinita var. asymmetrica, but I have not examined the specimen. As presently defined, the species is known from the type-locality (Easter Island), and the western Atlantic from North Carolina to Florida, the Gulf of Mexico, Argentina (Orensanz, 1974) and possibly from the Cape Verde Islands (Crossland, 1924) in depths from intertidal to 130 m. Arabella (Cenothrix) multidentata (Ehlers, 1887) Fig. 13 Aracoda multidentata Ehlers, 1887:112, pl. 34, figs. 8-10, pl. 35, figs. 1-4. Arabella maculosa.—Treadwell, 1921:114, pl. 9, figs. 12, 13, text-figs. 424— 428 [not Verrill, 1900]. Arabella iricolor.—Fauvel, 1923:438 [in part]—Hartman, 1938:12 [not Ne- reis iricolor Montagu, 1804]. Arabella novecrinita Crossland, 1924:71, text-figs. 89-95.—Monro, 1933a:260.—Gallardo, 1968:89, pl. 37, figs. 1-5. Arabella multidentata.—Hartman, 1938:12. Material examined.—FLORIDA: Blake Sta. 26, 24°37.5S'N, 83°36’W, 201 m, holotype (MCZ 825).—Hutchinson Island, Sta. II, 27°21.6'N, 80°13.2’W, 448 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 500 yum (i Toa Ei) / y' it Sie ees Soe 40 yum. ieeek eee — ety 20 ym ie ae Fig. 13. Arabella multidentata: a, Maxillae; b, Pygidium, dorsal view; c, d, Parapodia from middle segments, anterior view; e, Transversely serrated setae of middle segment; f, Lower winged capillary seta of middle segment; g, Asymmetrically hooded acicular setae from middle segments of Hutchinson Island specimen, anterior view; h, Same from holotype of Aracoda multidentata, anterior view. about 11 m, coarse calcareous sand; 1 specimen (USNM 54700).—Sta. V, 27°22.9'N, 80°13.9'W, about 11 m, coarse calcareous sand; 1 specimen (USNM 54699).—BIKINI: %4 mi W of SE point, 11-13 m; J. E. Morrison, col., 23 April 1946; 10+ specimens (USNM 23933, O. Hartman, det. as A. novecrinita). Description.—Holotype light yellowish-orange, iridescent; Hutchinson Island specimens red. Holotype relatively large, probably less than half complete, 33 mm long, 114 segments (Ehlers, 1887); Hutchinson Island spec- imens much smaller. Prostomium rounded, conical, with 4 dorsal posterior eyes. Peristomium of 2 apodus segments. Parapodia generally similar throughout body, sub-biramous, with about 2 fine notoacicula extending VOLUME 92, NUMBER 3 449 into lobed notopodial rudiment; setigerous lobe relatively short in anterior segments, longer posteriorly, with about 3 acicula often extending through integument in long, fine points; postsetal lobe about as long as setal lobe or slightly longer, extending nearly laterally from lower 74 of setal lobe. An- terior setae winged capillaries. Middle and posterior setae 4—5 of 4 general types: 1) upper seta winged capillary with wings smooth under high mag- nification; 2) seta below winged capillary with 6-7 prominent transverse rows of serrations (Fig. 13e); 3) winged capillary seta below transversely serrate seta shorter than upper winged capillary and with lightly serrate wings (Fig. 13f); 4) lower seta acicular, with relatively long, narrow, asym- metrical hood (Figs. 13g, h). Pygidium short, dorsally divided into 2 lobes, with 4 subequal anal cirri slightly longer than half width of pygidium. Mandibles anteriorly flared, fused along anterior half, widely divided and tapered posteriorly. Maxillae subsymmetrical: M I (1 + 6 — 7); M II (0 + 10 — 11); MITT (1 + 4 — 5); MIV(1 + 3 — 4); M V 1. Paired carriers about twice as long as maxillae; unpaired rod anteriorly rounded. Remarks.—The specimen from Tortugas, Florida, referred to Arabella novecrinita by Monro (1933a) is undoubtedly Arabella multidentata, the type-locality of which is near Tortugas in 201 m. Monro suggested the possible synonymy of A. novecrinita Crossland and A. multidentata (Eh- lers). He also suggested that A. maculosa Verrill, 1900, was the same, but the latter is a different species. Attempts by Fauvel (1923) and Hartman (1938) to synonymize A. multi- dentata with A. iricolor (Montagu) are incorrect, as the latter completely lacks acicular setae. The specimens from Easter Island, referred to A. mutans by Kohn and Lloyd (1973), were examined. Mouthparts are missing from one of the spec- imens (USNM 49534), apparently the one they described, but it appears to be A. multidentata. The other specimen (USNM 57434) is neither A. mutans nor A. multidentata but is apparently an undescribed species; formula for symmetrical maxillae of the latter specimen is: M I (2 + 7); M II (6 — 7); M III (1 + 5 — 6); MIV (1 + 3 — 4); and M V 1. Maxillae II of this spec- imen are symmetrical but are otherwise similar to those with 7 teeth on A. mutans (Chamberlin). Additionally, specimens reported as A. mutans by Day (1967:446) from southern Africa are probably A. multidentata. Based on references cited above, descriptions given for specimens incor- rectly referred to Arabella mutans (Chamberlin) by Kohn and Lloyd (1973), Gardiner (1976), and Day (1967), and material reported herein, A. multi- dentata is known from South Africa, Zanzibar, the Maldive Islands, Bikini Atoll, Easter Island and the east coast of North America from North Car- olina to Tortugas, Florida. Arabella multidentata has also been reported from southwest Australia and Amboina by Augener (vide Crossland, 1924). 450 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Arabella (Cenothrix) maculosa Verrill, 1900 Fig. 14 Arabella maculosa Verrill, 1900:651. Arabella novecrinita var. atlantica Crossland, 1924:78, text-figs. 96, 99-101, 105. Arabella iricolor.—Hartman, 1942b:55 [not Nereis iricolor Montagu, 1804]. Material examined.—BERMUDA: The Flats Inlet; A. E. Verrill, col., 1898; syntype (YPM 2716). Description.—Uncolored in alcohol. Body about 95 mm long, about 1 mm wide. Prostomium long, bluntly conical, with 4 posterior dorsal eyes, outer eyes larger. Peristomium of two relatively long apodus segments. Setal lobes relatively short, rounded, supported by 2 acicula in middle segments, with notopodial rudiment at upper edge supported by fine notoacicula; postsetal lobes subtriangular, about as long as setal lobes. Anterior setae winged capillaries; middle and posterior parapodia with 4—5 setae of 4 general types: 1) upper seta long, geniculate, winged capillary, finely denticulate proxi- mally (Fig. 14c); 2) shorter seta below winged capillary with 5—6 prominent transverse rows of serrations proximally and fine denticles laterally (Fig. 14d); 3) setae below transversely serrate seta similar to upper seta but short- er, with longer region of slightly more prominent denticles (Fig. 14e); and 4) lower, asymmetrically hooded, acicular seta (Fig. 14f). Pygidium with indistinct pair of lateral, rounded lobes without anal cirri (Fig. 14a). Maxillae II asymmetrical: MI (1 + 6 — 7); MII (6 — 8) + (1 + 10 — 12); MII(1 + 3 —-5);MIV(1_+3-4);MV1. Remarks.—Hoods of acicular setae of A. maculosa are much more ob- viously asymmetrical than are those of A. multidentata and A. mutans. These acicular setae were apparently overlooked by Hartman (1942b), who attempted to combine the species with A. iricolor (Montagu). The original description of A. maculosa was brief and lacked most important characters. My examination of Verrill’s syntype indicates no apparent differences from A. novecrinita var. atlantica described by Crossland (1924) from the Cape Verde Islands. A. novecrinita var. logani Crossland, 1924, from Suez, ap- parently differs in having parapodia with a larger number of setae and aci- cula. Crossland’s text-fig. 96, attributed to be maxillae of A. novecrinita var. logani, apparently was actually of A. novecrinita var. atlantica, be- cause he indicated that the specimen from which it was drawn was from Cape Verde Islands. The species is known only from localities indicated above. Family Dorvilleidae Chamberlin, 1919 I am following the revision of this family by Jumars (1974), with one exception: Papilliodorvillea Pettibone, 1961 is retained rather than referred VOLUME 92, NUMBER 3 451 Fig. 14. Arabella maculosa (syntype, YPM 2716): a, Posterior end, lateral view; b, Para- podium from middle segment, anterior view; c, Upper winged capillary seta of middle para- podium; d, Transversely serrated setae of same; e, Lower winged capillary setae from same; f, Asymmetrically hooded acicular setae from same, anterior view. (a, b, d, and e not scaled). to Dorvillea. Nomenclature of mouthparts follows Jumars. Free denticles are numbered sequentially from basal plates. Genus Dorvillea Parfitt, 1866 Dorvillea sociabilis (Webster, 1879) Fig. 15 Staurocephalus sociabilis Webster, 1879:243, pl. 7, figs. 89-91. Dorvillea sociabilis.—Hartman, 1945:27, pl. 5, figs. 1, 4, 5; 1951:66, pl. 8, figs. 3, 5.—Rullier, 1974:52.—Gardiner, 1976:215, figs. 29]—n. Material examined.—FLORIDA: Hutchinson Island, Sta. II, 27°21.6'N, 80°13.2’W, about 11 m, coarse calcareous sand; | specimen (FSBC I 18873).—Sta. IV, 27°20.7’N, 80°12.8’W, about 11 m, coarse calcareous sand; 3 specimens (FSBC I 18870, 18871).—Sta. V, 27°22.9’N, 80°13.9’W, about 11 m, coarse calcareous sand; 3 specimens (USNM 54705, 54706; FSBC I 18872). 452 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON =e 20 um, Fig. 15. Dorvillea sociabilis: a, Fused maxillary carriers, basal plates and posterior free denticles (D 1-D 4) of right upper row of maxillae; b, Fused upper maxillary carriers, dorsal view; c, Right basal plates of upper and lower rows of maxillae; d, Left basal plate and posterior free denticles (D 1—D 3) of lower row of maxillae; e-l, Free denticles from upper row of maxillae (total number 18): e, D 1; f, D 2; g, D 3—D 5; h, D 9; i, D 10; j, D 15; k, D 17; 1, D 18; m-p, Free denticles from lower row of maxillae (total number 26): m, Middle denticle (D 13); n, D 21; 0, D 24; p, D 26; q, Tip of long simple seta; r, Compound falciger (q, r not scaled). Description.—Maxillae with pair of carriers and 4 rows of denticles with accompanying basal plates. Upper carriers fused, with posterior knob and 2 anterior wing-like pieces attached to basal plates of upper maxillary rows; wings minutely serrate on attachment border; lower carriers small, fused medially to basal plates of upper and lower rows of maxillae. Upper rows with stout subequal basal plates, with 9 teeth on left and 10 on right, with 17-18 free denticles; first (D 1) with 3 teeth, stouter main tooth, smaller lateral and medial teeth (Fig. 1Se); D 5 with stout recurved tooth, 1 lateral and 3 medial (Fig. 15g); D 15 with long, sharply pointed main tooth, | lateral and 2 medial (Fig. 15j); anterior denticles slender, hollow, with 1—2 points (Figs. 15k, 1). Lower row of maxillae with asymmetrical basal plates, shorter right one with 7 teeth, longer left one with 10 teeth; proportional lengths of left and right basal plates 3:2 (Figs. 15c, d); proportional lengths of right lower and upper basal plates 2:5; 23-27 free denticles anterior to lower basal plates (Figs. 15d, m—p); D 1 with 1 main tooth, | lateral, 2 medial; D 3 with main tooth, 1 lateral, 3 medial; D 14 with 2 main teeth of similar size and 2 small (Fig. 15m); D 23 with 2 large teeth and 1 small (Fig. 15n); D 24 with VOLUME 92, NUMBER 3 453 2 equal teeth (Fig. 150); anterior denticle (D 26) with 1 tooth (Fig. 15p); anterior teeth (about D 20-26) long, slender, hollow. Teeth of basal plates of upper and lower rows increasing in size anteriorly; those of upper row much larger than lower. One specimen with maxillae completely duplicated (i.e., 8 rows of maxillae), indicating that maxillae are replicated. Supra- acicular simple setae of 2 types: 1) long, slender, finely serrate on basal half; and 2) short, stout, serrate up to tip; both flattened, with bifid tips (Fig. 15q). Subacicular compound hooks with distal ends of shafts strongly toothed on longest part; blades with toothed edges and bidentate tips (Fig. i510). Remarks.—Dorvillea sociabilis was described from Virginia, and is pres- ently known from Virginia to Hutchinson Island, Florida, Cuba (Rullier, 1974) and the eastern Gulf of Mexico in 0-160 m. Genus Protodorvillea Pettibone, 1961 Protodorvillea bifida, new species Figs. 16, 17a—f Material examined.—FLORIDA: Hutchinson Island, Sta. IV, 27°20.7’N, 80°12.8’W, about 11 m, coarse calcareous sand; holotype (USNM 57471, R. Gallagher, col., March 1972); 4 paratypes (AHF POLY 1272; FSBC I 20474- 20476).—Sta. II, 27°21.6’N, 80°13.2'W, about 11 m, coarse calcareous sand, 9 paratypes (AHF POLY 1271; ZMH P-15540; FSBC I 20467-—20473).—Sta. V, 27°22.9'N, 80°13.9'W, about 11 m, coarse calcareous sand, 3 paratypes (USNM 57472; FSBC I 20477).—Lower Tampa Bay, 27°36'15’N, 82°43'22"”W, 8 m, sand; J. Taylor and C. Saloman, cols., 29 October 1963; 3 paratypes (USNM 57473; ZMH P-15541; FSBC I 20478). Description.—Colorless in alcohol; eyes red. Largest specimen about 10 mm long, 0.25 mm wide, about 80 segments. Prostomium about as long as wide, generally flattened dorsoventrally, bluntly conical anteriorly, slightly convex laterally, slightly convex to concave posteriorly. Antennae origi- nating on posterior half of prostomium, about half as long as greatest pros- tomial width, usually biarticulate. Palps originating laterally, slightly ante- rior to middle of prostomium, about equal in length to first 10 segments. Eyes usually 2 pairs; posterior pair larger, with lens, anterior and slightly lateral to origins of antennae; anterior pair much smaller, apparently sub- dermal, anterior and medial to origins of palps, not visible in mature spec- imens (Figs. 16a, b). Peristomium of 2 apodus segments, each slightly nar- rower and about as long as following segments. Parapodia uniramous, without notoacicula, supported by single neuroaciculum, subcylindrical, with digitiform dorsal and ventral cirri near tips, with indistinct presetal and postsetal lobes, with subacicular setal lobe often more elongate than supra- acicular lobe. Setae of 6 types: 1) 1-2 upper simple serrate capillaries; 2) 454 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 16. Protodorvillea bifida: a, Anterior end of mature specimen, dorsal view; b, Anterior end of immature specimen, dorsal view; c, Posterior end, ventrolateral view; d, Mandibles; e, Maxillae. furcate seta; 3) cultriform seta replacing furcate seta in few posterior seg- ments; 4) subacicular compound falcigers with bidentate tips, 4—5 in anterior parapodia, 2-3 in posterior; 5) lower simple seta in posterior few segments; 6) 2-6 natatory often originating posterior to compound falcigers in gravid VOLUME 92, NUMBER 3 455 segments of sexually mature specimens. Furcate setae smooth-sided, with tips of unequal length; longer tine slightly winged, with papilliform tip; short- er tine not winged, with bifid tip. Upper falcigerous blades longer, lower blades shorter; greatest length about 50 um; bidentate tips with secondary tooth subequal to primary tooth. Cultriform setae serrate, with short, stout subequal teeth, bidentate tips similar to those of falcigers. Pygidium some- what obliquely truncate, with 2 pairs of anal cirri, long, filiform, non-artic- ulated dorsal pair and short, stout ventral pair. Sexually mature specimens with sex products beginning in setigers 10-13, continuing to near posterior end. Mandibles (Fig. 16d) anteriorly flared, with 4—S fused and 3-5 free teeth on each side; inner teeth with small denticles on elongate median edge; mandibles fused posterior to denticles of inner teeth, posteriorly elongate. Maxillae with pair of carriers, 2 pairs of basal plates, 2 paired rows of free denticles continuous with basal plates (Fig. 16e); upper carriers toothed on anterior dorsal parts, adnate to posterior halves of basal plates of upper row of maxillae, ending posteriorly within brownish area of maxillary ligament; lower carriers appearing as reinforced area of maxillary membrane; ligament in area of carriers only slightly thickened. Basal plates of upper row rela- tively massive, with 4—S main teeth, about twice as many secondary teeth; teeth decreasing in size posteriorly; 12-13 free denticles; D 1 very stout, with 2 teeth, large lateral, small medial; D 2 with 5—6 teeth, 1-2 small medial ones, | large hook, 2 small hooks, 1 large lateral tooth; next 4 denticles with additional small lateral tooth, larger teeth becoming smaller; denticles on anterior end with teeth gradually reduced in number, becoming subequal in size, less strongly hooked; anterior denticle of one side with 2 teeth, that of opposite side with 4; free denticles of upper row stout, thickened. Lower row of maxillae with thin, narrow, asymmetrical basal plates with about Same number and arrangement of teeth as upper row; teeth smaller, more sharply pointed; 19 free denticles on each side; D 1 with about seven small, acutely pointed, posteriorly directed teeth, middle tooth appearing slightly larger; thereafter, teeth of denticles becoming more numerous, smaller, gradually disappearing except median tooth which continues anteriorly, pos- sibly absent from anteriormost 1-2 denticles; free denticles of lower row thin, flattened. Remarks.—Furcate setae with one bifid tine have not been noted previ- ously for species of Protodorvillea. The bidentate blades of the compound falcigers with subequal teeth differ from P. kefersteini, in which the blades show a minute secondary tooth. To my knowledge the upper cultriform and lower simple setae observed on P. bifida and P. kefersteini from Hutchinson Island have not been previously reported for Protodorvillea. Etymology.—The specific name is derived from the Latin and refers to the single bifid tine of the furcate setae. 456 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Protodorvillea kefersteini (McIntosh, 1869) Figs. 17g—m Staurocephalus kefersteini McIntosh, 1869:417, pl. 16, fig. 11. Protodorvillea kefersteini.—Pettibone, 1961:179.—Hartman-Schroder, 1971:262, figs. 87a-g.—Hobson, 1971:542, fig. 8.—Orensanz, 1973a:335, pl. 4, figs. 1-9.—Gardiner, 1976:214, figs. 29h—-k. Material examined.—FLORIDA: Hutchinson Island, Sta. IV, 27°20.7'N, 80°12.8’W, about 11 m, coarse calcareous sand; 1 specimen (FSBC I 18874).—Sta. V, 27°22.9’N, 80°13.9’W, about 11 m, coarse calcareous sand; 6 specimens (USNM 54707-54709; FSBC I 18875, 18876).—MASSACHU- SETTS: Cape Cod Bay, 15.5 m, gravelly sand; D. C. Grant, col., Kay Hobson, det.; 8 specimens (USNM 42027). Description.—Prostomium with 2 pairs of eyes on most specimens; an- terior pair medial to origins of palps; posterior pair slightly larger, medial to origins of antennae. Blades of compound falcigers (Figs. 17j, k) with secondary tooth much smaller than primary, as figured by Hartman-Schro- der (1971:fig. 87e). Forked setae with tines slightly asymmetrical in size and shape. Modified simple, cultriform setae (Fig. 171) replacing forked setae in posterior few segments, moderately curved near tip, with teeth on convex border longer basally, shorter near tip; simple setae (Fig. 17m) smooth, slender, with slightly curved, unidentate tips in lower part of posterior few parapodia. Mouthparts as figured by Orensanz (1973a:pl. 4, figs. 3, 4) and Hartman- Schroder (1971:figs. 87f, g). Maxillae replicated in 1 of 2 specimens dis- sected. Remarks.—Protodorvillea kefersteini was described originally from Scot- land and is presently known from Ireland, the English Channel and North Sea, Mediterranean Sea, Black Sea, Argentina and Atlantic coast of North America from Massachusetts to Florida. Cultriform setae are absent from New England specimens examined. Genus Schistomeringos Jumars, 1974 Schistomeringos pectinata, new species Figs. 18-20 Material examined.—FLORIDA: Hutchinson Island, Sta. IV, 27°20.7'N, 80°12.8’'W, about 11 m, coarse calcareous sand; holotype (USNM 57474, R. Gallagher, col., July 1972), 9 paratypes (ZMH P-15543; FSBC I 20496- 20503).—Sta. II, 27°21.6'N, 80°13.2’W, about 11 m, coarse calcareous sand; 10 paratypes (USNM 57475; AHF POLY 1273, 1274; FSBC I 20484- 20494).—Sta. III, 27°22.0’N, 80°12.4’W, about 7 m, medium calcareous sand; 1 paratype (FSBC I 20495).—Sta. V, 27°22.9'N, 80°13.9’W, about 11 VOLUME 92, NUMBER 3 457 50 um [ica ee 10 um k 3 50 ym 10 ym Fig. 17. Protodorvillea bifida: a, Parapodium of middle segment, anterior view; b, Forked seta from same; c, Upper compound falciger from same; d, Middle compound falciger from same; e, Lower simple seta; f, Cultriform seta. Protodorvillea kefersteini: g, Anterior end, dorsal view; h, Pygidium, ventral view; i, Forked seta; j, Upper compound falciger; k, Lower compound falciger; 1, Cultriform seta; m, Lower simple seta. m, coarse calcareous sand; 2 paratypes (ZMH P-15542; FSBC I 20504).— Lower Tampa Bay, J. Taylor and C. Saloman, cols., Oct. 1963; 1 paratype (USNM 57476). Description.—Colorless in alcohol; eyes red. Largest specimen about 15 458 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 10 um Fig. 18. Schistomeringos pectinata: a, Anterior end (holotype), dorsal view; b, Anterior end of immature specimen, dorsal view; c, Same, lateral view; d, Posterior end, dorsal view; e, Maxillary basal plates and carriers of upper and lower rows of maxillae; f-k, Free denticles of upper row of maxillae (total number 25): f, D 1; g, D 3; h, D 7; i, D 21; j, D 24; k-n, Free denticles of lower row of maxillae (total number 30): k, D 1; 1, D 4; m, D 10; n, D 28. VOLUME 92, NUMBER 3 459 50 um Fig. 19. Schistomeringos pectinata: a, Parapodium of setiger 1 of holotype, posterior view; b, Parapodium of setiger 12 of immature specimen, anterior view; c, Parapodium of setiger 20 of immature specimen, anterior view; d, Same of holotype, posterior view; e, Parapodium of middle segment of immature specimen, anterior view; f, Parapodium of middle segment, sub- acicular setal lobe partially extended, anterior view. mm long, slender, about 90 segments. Prostomium flattened, blunily conical; generally straight posteriorly, sides slightly concave. Antennae originating dorsolaterally and slightly posterior to middle of prostomium, equal in length to palps when fully developed, with 8-9 articles. Palps originating laterally and slightly anterior to antennae, biarticulate, with long, stout, tapered, generally wrinkled palpophores and elongate, oval palpostyles. Eyes of ma- 460 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON we? aA 5 be Re ime oa Oe eon i. Tee / 1O um i Fig. 20. Schistomeringos pectinata: a, Parapodium of middle segment, posterior view; b, Forked setae, opposite views (holotype); c, 3 compound setae of setiger 20 (holotype). ture specimens one pair, enlarged, reniform, covering almost total area be- tween palps and antennae (Fig. 18a); eyes of immature specimens 2 pairs (Figs. 18b, c) or none; larger pair slightly anterolateral to antennae; smaller pair medial to palps (Fig. 18c). Peristomium of 2 apodus segments, anterior one often slightly longer, narrower than following segments. Parapodia of setiger | (Fig. 19a) uniramous, without dorsal cirri; remainder sub-biramous, with slender notoaciculum extending into cirrophores of dorsal cirri. Dorsal cirri generally longer than setal lobes in anterior segments, slightly shorter posteriorly; proportional lengths of cirrophores and cirrostyles varying from slightly greater than 1:1 to slightly less than 2:1; separation of cirrophores and cirrostyles often indistinct (Fig. 20a). Setal lobes with long, slender aciculum in upper part; supra-acicular setal lobe rounded, dorsally broad; subacicular setal lobe in anterior segments separated from supra-acicular lobe, broadly rounded in anterior view; distinct presetal and postsetal lobes extending from above supra-acicular lobe and ending at ventral cirrus; pre- VOLUME 92, NUMBER 3 461 setal lobe less developed than postsetal; subacicular setal lobes in middle and posterior segments (Figs. 19d, 20a), extending from ventral edge as long, thin triangular processes, with obscure presetal lobe in similar position as in anterior segments and more prominent postsetal lobe. Immature spec- imens with supra-acicular setal lobe longest and subacicular setal lobe not extended, with short presetal lobe and slightly longer postsetal lobe both visible in anterior view (Figs. 19b, c, e); specimen with partially extended subacicular setal lobe figured (Fig. 19f). Ventral cirri originating subdistally on parapodial lobes, usually extending about to tips. Single aciculum col- orless; tip abruptly narrowed, acute. Supra-acicular setae of 2 types: 1) 1- 2 forked setae, and 2) 1-6 (usually 2-3) capillaries. Subacicular setae 6-10 compound falcigers. Forked setae with slightly unequal tines; shorter tine cusped (Fig. 20b), with 4 cusps on one side, 3 extending to edge, 2 extending to opposite side; shorter tine faintly limbate above cusps. Capillary setae serrate on one edge. Blades of compound falcigers serrate; lengths of blades varying between 18 and 64 um, longer ones dorsal; tips bidentate, hooded; shafts of falcigers with 3 serrations below tip of long extension. Supra-aci- cular cultriform setae on parapodia of first setiger of Tampa Bay specimen (USNM 57476). All’ segments with 2 transverse dorsal rows of cilia con- tinuing on dorsal and ventral sides of cirrophores of dorsal cirri and para- podial lobes. Pygidium (Fig. 18d) rounded, obliquely truncate, with 2 pairs of anal cirri, long filiform dorsal pair and stouter ventral pair. Mandibles with 3-6 fused and 3-5 free teeth on each side. Maxillary supports as figured (Fig. 18e). Basal plates of upper row of maxillary struc- tures with 7 principal teeth, 17-20 secondary teeth; up to 27 free denticles, usually 20-25; D 1 with 4 teeth, 1 principal tooth, 2 lateral and 1 medial teeth (Fig. 18f); free denticles thereafter gradually changing anteriorly, first having 2 principal teeth with smaller teeth, 3 principal teeth with smaller teeth and nearly subequal teeth near anterior end; D 3 with 7 teeth, 2 large, 5 smaller (Fig. 18g); D 7 with 10 teeth, 3 large, 7 smaller (Fig. 18k); D 21 with 10 rounded teeth, lateral and medial ones somewhat separated from others, falcate except for lateral one (Fig. 181); D 24 with about 10 subequal, rounded teeth. Basal plates of lower row with 10-11 principal teeth, about 25 secondary teeth, plates asymmetrical in length in some specimens and 14-*% as long as those of upper row with smaller teeth; up to 37 free den- ticles, usually about 30. On lower row, D 1 with 1 larger tooth, 3-5 medium teeth, several smaller teeth (Fig. 18k); D 4 with 9 large and 10 small teeth, with medial tooth pointed obliquely toward midline, others hooked poste- riorly; denticles beginning with about D 7 or more anteriorly, with medially directed hook and numerous small, posteriorly curved hooks on nearly straight edge (Figs. 18m, n); teeth on anterior denticles easily broken, often absent. Maxillae in various stages of replication in more than 10% of spec- imens. 462 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Remarks.—Species of Schistomeringos were summarized by Jumars (1974:104) and Westheide (1977:380). Adults of S. pectinata resemble S. neglecta (Fauvel, 1923:447, figs. 179i-q) in having one pair of eyes and parapodia with an extended subacicular neuropodial lobe. However, the forked setae have no serrations below the shorter tine. Mouthparts of S. neglecta were insufficiently described to allow comparison with S. pectin- ata. S. pectinata resembles S. pacifica (Westheide, 1977:379) from the Gala- pagos Islands in having similar mouthparts and parapodia. The latter species lacks eyes and has well defined nuchal organs on the anterior border of the first segment, and stout, bidentate-tipped supra-acicular, simple cultriform setae on posterior parapodia. S. mossambica (Hartman-Schroder, 1974:30) from Mozambique has a similar prostomium, eyes and parapodia to S. pectinata. However, the par- apodia of S. mossambica are much shorter compared to body width than are those of S. pectinata. Details of maxillae and presence of dorsal ciliary bands were not described for S. mossambica. Etymology.—The specific name, derived from the Latin, means comb- shaped and refers to shape of the free denticles of Maxillae II. Acknowledgments The following individuals loaned specimens: Drs. Marian H. Pettibone and Meredith L. Jones, National Museum of Natural History; Dr. Willard D. Hartman, Peabody Museum of Natural History; Dr. Herbert W. Levi, Museum of Comparative Zoology; Mr. Roy Eolerdd, Naturhistoriska Riks- museet; Dr. G. Hartwich, Zoologisches Museum, Berlin; and Drs. J. D. George and A. I. Muir, British Museum (Natural History). Dr. Norman J. Blake and Mr. Robert G. Ernest, Department of Marine Science, University of South Florida, donated specimens. Dr. Pettibone, Dr. Kristian Fauchald, Allan Hancock Foundation, and Dr. Joseph L. Simon, University of South Florida, commented on early drafts of the manuscript and provided infor- mation, and Dr. Pettibone thoroughly reviewed the final draft. Dr. Pettibone and Dr. Jones courteously provided space and assistance during a visit to the Smithsonian Institution. Mr. James H. Seagle of the Marine Research Laboratory assisted with some figures. Many other individuals from the Marine Research Laboratory and Applied Biology, Inc., Jensen Beach, Florida, participated in planning and execution of the Hutchinson Island study for which Florida Power and Light Co., Inc. provided partial funding. All are gratefully acknowledged. Literature Cited Augener, H. 1922. Ueber litorale Polychaeten von Westindien.—Ges. Naturf. Freunde Berlin, Sitzber. 1922:38-63. VOLUME 92, NUMBER 3 463 Chamberlin, R. V. 1919. The Annelida Polychaeta.x—Mem. Mus. Comp. Zool. Harv. Univ. 48. 514 pp., 80 pls. Crossland, C. 1924. Polychaeta of tropical East Africa, the Red Sea and Cape Verde Islands, collected by Cyril Crossland and of the Maldive Archipelago collected by Professor Stanley Gardiner, M.A., F.R.S.: The Lumbriconereidae and Staurocephalidae.—Proc. Zool. Soc. Lond. 1924:1—-106. Day, J. H. 1963. The polychaete fauna of South Africa. Part 8: new species and records from grab samples and dredgings.—Bull. British Mus. (Nat. Hist.) Zool. 10:381-445. . 1967. A monograph on the Polychaeta of southern Africa. Part 1.—Br. Mus. (Nat. Hist.) Publ. No. 656:458 pp. . 1973. New Polychaeta from Beaufort, with a key to all species recorded from North Carolina.—NOAA (Natl. Ocean. Atmos. Adm.) Tech. Rep. NMFS (Natl. Mar. Fish. Serv.) Circ. 375. 140 pp. Ehlers, E. 1887. Report on the annelids of the dredging expedition of the U.S. Coast Survey steamer Blake.—Mem. Mus. Comp. Zool. Harv. Univ. 15, vi + 335 pp., 60 pls. Fauchald, K. 1977. Polychaetes from intertidal areas in Panama, with a review of previous shallow-water records.—Smithson. Contrib. Zool. No. 221, 81 pp. Fauvel, P. 1923. Polychétes errantes.—Faune de France 5, 488 pp. Gallagher, R. M. 1977. Nearshore marine ecology at Hutchinson Island, Florida: 1971-1974. II. Sediments.—Florida Mar. Res. Publ. No. 23:6—24. , and M. L. Hollinger. 1977. Nearshore marine ecology at Hutchinson Island, Florida: 1971-1974. I. Introduction and rationale: 1971—1975.—Florida Mar. Res. Publ. No. 23:1-S. Gallardo, V. A. 1968. Polychaeta from the Bay of Nha Trang, South Viet Nam.—NAGA Rep. [1967] 4(3):35—279. — Gardiner, S. L. 1976. Errant polychaete annelids from North Carolina.—Jour. Elisha Mitchell Sci. Soc. [Fall, 1975] 91(3):77-220. Hall, J. R., and C. H. Saloman. 1975. Distribution and abundance of macroinvertebrate species of six phyla in Tampa Bay, Florida, 1963-64 and 1969.—NMES Data Rep. 100. 505 pp. Hartman, O. 1938. Annotated list of the types of polychaetous annelids in the Museum of Comparative Zoology.—Bull. Mus. Comp. Zool., Harv. Univ. 85:1-31, 3 pls. 1942a. The identity of some marine annelid worms in the United States National Museum.—Proc. U.S. Natl. Mus. 92:101-140. . 1942b. A review of the types of polychaetous annelids at the Peabody Museum of Natural History, Yale University.—Bull. Bingham Oceanogr. Collect. Yale Univ. 8(1): 1-98. 1944a. Polychaetous annelids. Part 5. Eunicea.—Allan Hancock Pacific Exped. 10(1): 1-238. 1944b. New England Annelida. Part 2, including the unpublished plates by Verrill with reconstructed captions.—Bull. American Mus. Nat. Hist. 82:327-343. . 1945. The marine annelids of North Carolina.—Bull. Duke Univ. Mar. Sta. 2:1—54. . 1948. The marine annelids erected by Kinberg with notes on some other types in the Swedish State Museum.—Ark. Zool. 42A(1): 1-137. . 1951. The littoral marine annelids of the Gulf of Mexico.—Publ. Inst. Mar. Sci. Univ. Texas 2:7-124. 1959. Catalogue of the polychaetous annelids of the world, parts 1 and 2.—Allan Hancock Found. Publ. Occas. Pap. 23:1-628. . 1965. Deep water benthic polychaetous annelids off New England to Bermuda and other North Atlantic areas —Allan Hancock Found. Publ. Occas. Pap. 28:1—378. Hartman-Schroder, G. 1971. Annelida, Borstenwiirmer, Polychaeta.—Tierwelt Dtl. 58:1-594. . 1974. Weitere Polychaeten von Ostafrika (Mogambique und Tansania).—Mitt. Ham- burg Zool. Mus. Inst. 71:23-33. 464 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Hobson, K. D. 1971. Some polychaetes of the Superfamily Eunicea from the North Pacific and North Atlantic Oceans.—Proc. Biol. Soc. Washington 83(47):527—544. Imajima, M., and M. Higuchi. 1975. Lumbrineridae of polychaetous annelids from Japan, with descriptions of six new species.—Bull. Natl. Sci. Mus. Tokyo, Ser. A. 1(1):5-37. Jumars, P. A. 1974. A generic revision of the Dorvilleidae (Polychaeta), with six new species from the deep North Pacific.—Zool. Jour. Linn. Soc. Lond. 54:101-135. Kinberg, J. G. H. 1865. Annulata nova.—Ofvers. K. Vet. Akad. Forh. Stockholm 21:559- 574. Kohn, A. J., and M. C. Lloyd. 1973. Marine polychaete annelids of Easter Island.—Int. Rev. Gestamten Hydrobiol. 58(5):691-712. McIntosh, W. C. 1869. On the structure of the British nemerteans, and some new British annelids.—Trans. Roy. Soc. Edinburgh 25:30S—433. Monro, C. C. A. 1933a. On a collection of Polychaeta from Dry Tortugas, Florida.—Ann. Mag. Nat. Hist. London, Ser. 10, 12:244—-269. . 1933b. The Polychaeta Errantia collected by Dr. C. Crossland at Colon, in the Panama region, and the Galapagos Islands during the expedition of the S.Y. St. George.—Proc. Zool. Soc. London 1933(1): 1-96. Moore, J. P. 1904. New Polychaeta from California.—Proc. Acad. Nat. Sci. Philadelphia 56:484—-503, pls. 37, 38. . 1909. The polychaetous annelids dredged in 1908 by Mr. Owen Bryant off the coasts of Labrador, Newfoundland and Nova Scotia.—Proc. U.S. Natl. Mus. 37(1703): 133-146. . 1911. The polychaetous annelids dredged by the U.S.S. ‘‘Albatross’’ off the coast of southern California in 1904: III. Euphrosynidae to Goniadidae.—Proc. Acad. Nat. Sci. Philadelphia 63:234-318, pls. 15-21. Orensanz, J. M. 1973a. Los anélidos poliquetos de la Provincia Biogeografica Argentina. III. Dorvilleidae.—Physis Secc. A Océanos Org. 32(85):325—342. . 1973b. Los anélidos poliquetos de la Provincia Biogeografica Argentina. IV. Lum- brineridae.—Physis Secc. A Océanos Org. 32(85):343-393. . 1974. Los anélidos poliquetos de la Provincia Biogeografica Argentina. VI. Arabel- lidae.—Physis Secc. A Océanos Org. 33(87):381—408. Pettibone, M. H. 1961. New species of polychaetes from the Atlantic Ocean, with a revision of the Dorvilleidae.—Proc. Biol. Soc. Washington 74: 167-186. . 1963. Marine polychaete worms of the New England region. 1. Aphroditidae through Trochochaetidae.—Bull. U.S. Natl. Mus. 227(1):1-356. Ramos, J. M. 1976. Lumbrineridae (Polychéte Errantes) de Méditerranée.—Ann. Inst. Ocean- ogr. Paris 52(1):103—137. Rullier, F. 1974. Quelques annelides polycheétes de Cuba recueillies dans des eponges.—Trav. Mus. Hist. Nat. ‘“‘Grigore Antipa’’ 14:9-77. Taylor, J. L. 1971. Polychaetous annelids and benthic environments in Tampa Bay, Florida.— Ph.D. Dissertation, Univ. Fla., Gainesville, Fla. 1332 pp. Treadwell, A. L. 1921. Leodicidae of the West Indian region.—Carnegie Inst. Washington Publ. No. 293, Pap. Tortugas Lab. 15, 131 pp. 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—Rep. U.S. Comm. Fish Fish. for 1871—1872:295-778. . 1874. Explorations of Casco Bay by the U.S. Fish Commission, in 1873.—Proc. Am. Assoc. Adv. Sci. 22B:340-395, 6 pls. . 1875. Brief contributions to zoology from the Museum of Yale College. Results of dredging expeditions off the New England coast in 1874.—American Jour. Sci. Arts 10:36-43, pls. 3, 4. . 1879. Notice of recent additions to the marine invertebrata of the northeastern coast of America, with descriptions of new genera and species and critical remarks on others. VOLUME 92, NUMBER 3 465 Part I. Annelida, Gephyraea, Nemertina, Nematoda, Polyzoa, Tunicata, Mollusca, An- thozoa, Echinodermata, Porifera.—Proc. U.S. Natl. Mus. 2:165-—20S. . 1900. Additions to the Turbellaria, Nemertina, and Annelida of the Bermudas, with revisions of some New England genera and species.—Trans. Conn. Acad. Arts Sci. 10:595—671, 1 pl. Webster, H. E. 1879. Annelida Chaetopoda of the Virginia coast.—Trans. Albany Inst. 9:202- 269. , and J. E. Benedict. 1887. The Annelida Chaetopoda from Eastport, Maine.—Rep. U.S. Fish Comm. for 1885:707-755S, 8 pls. Westheide, W. 1977. Interstitielle fauna von Galapagos. XVIII. Nereidae, Eunicidae, Dor- villeidae (Polychaeta).—Mikrofauna Meeresboden 63:357-393. Worth, D. F., and M. L. Hollinger. 1977. Nearshore marine ecology at Hutchinson Island, Florida: 1971-1974. HII. Physical and chemical environment.—Fla. Mar. Res. Publ. No. 23:25-85. Marine Research Laboratory, 100 ooo Avenue S.E., St. Petersburg, Florida 33701. PROC. BIOL. SOC. WASH. 92(3), 1979, pp. 466-481 FOUR NEW SPECIES OF CARAZZIELLA (POLYCHAETA: SPIONIDAE) FROM NORTH AND SOUTH AMERICA, WITH A REDESCRIPTION OF TWO PREVIOUSLY DESCRIBED FORMS James A. Blake Abstract.—Four new species of Carazziella are described from North and South America: C. hobsonae from Florida and Texas, C. calafia from cen- tral California, C. carrascoi from Chile and C. patagonica from Argentina. Carazziella citrona (Hartman, 1941), the type-species, is redescribed from the California type-collection and C. reishi (Woodwick, 1964) from the Mar- shall Islands is redescribed from the holotype and from new materials from Indonesia and Johnston Atoll. Relationships of the American species with counterparts in Australasia are discussed along with a consideration of the peculiar gizzard-like structure found in some species of Carazziella and Polydora. The genus Carazziella was recently proposed by Blake & Kudenov (1978) in a study of Australian Spionidae. In their paper, 4 new species from Aus- tralia are described and 3 previously known species are referred to the genus: C. citrona (Hartman, 1941), designated as the type-species, C. reishi (Woodwick, 1964) from the Marshall Islands and C. quadricirrata (Rainer, 1973) from New Zealand. Carazziella belongs to the so-called Polydora-complex and is distin- guished from the other genera by having 2 types of major spines on the modified fifth setiger and by lacking branchiae anterior to setiger 5. Nine of the 11 species now known, including those described as new in this paper, have bristles on both types of the major spines. Four new species of Carazziella from North and South America are de- scribed herein. The type-species, C. citrona (Hartman, 1941) from Califor- nia, has been reexamined and is redescribed. The holotype of C. reishi (Woodwick, 1964) from Enewetak is redescribed and compared with addi- tional materials from Indonesia and Johnston Atoll. Thanks are extended to the following individuals for the loan of materials upon which this report is based: Dr. James Nybakken, Moss Landing Marine Laboratories, Moss Landing, California; Dr. John L. Taylor, then of the National Marine Fish- eries Service, St. Petersburg Beach, Florida; Mr. Clyde A. Henry, Texas A&M University, Galveston, Texas; Ms. Linda Ward, Naval Oceans Sys- tems Center (NOSC), Kailua, Hawaii; Dr. José M. Orensanz, Instituto de VOLUME 92, NUMBER 3 467 Biologia Marina, Mar del Plata, Argentina; Dr. Franklin D. Carrasco, In- stituto de Biologia, Concepcion, Chile; Dr. Kristian Fauchald, Allan Han- cock Foundation (AHF) and Dr. Marian H. Pettibone, National Museum of Natural History, Smithsonian Institution (USNM). Mrs. Karen Green and Mr. Bob Osborn assisted my visit to AHF in May, 1978 by providing spec- imens and data. Mr. William J. Light read and commented on the manu- script. Type-materials of the newly described species are deposited in the Smith- sonian Institution (USNM), California Academy of Sciences (CAS), Museo Argentino de Ciencias Naturales, Buenos Aires (MACN) and Museo Zool- ogico, Instituto de Biologia, Universidad de Concepcion, Chile (MZC). Genus Carazziella Blake & Kudenov, 1978 Type-species.—Polydora citrona Hartman, 1941, designated by Blake & Kudenov. Diagnosis.—Prostomium rounded or incised, extending posteriorly as a caruncle; eyes present. Setiger | with or without notosetae. Setiger 5 mod- ified, with 2 types of heavy spines arranged in a double curved row: (1) first type with expanded tip bearing bristles or cusps; (2) second type simple, falcate; both types usually bristle-topped; dorsal fascicles of capillaries pres- ent or absent; with a well-developed ventral fascicle of capillaries. Bidentate neuropodial hooded hooks without constriction on shaft, beginning on se- tiger 7-14, with conspicuous angle between teeth, both teeth forming oblique angle with shaft. Branchiae beginning posterior to setiger 5. Pygi- dium with 2-4 lobes or fingerlike cirri. Carazziella citrona (Hartman, 1941) emended Fig. 1 Polydora citrona Hartman, 1941:311-312, pl. 45, figs. 9-16; 1961:29; 1969: 131-132. [Not Carrasco, 1974:193; 1976:25—28 (see below)]. Material examined.—CALIFORNIA: Mission Bay, Sept, 1938, coll. Olga Hartman.—holotype (AHF Poly 33) and 3 paratypes (AHF Poly 1266). Description.—A large species, up to 30 mm long and 1.0 mm wide for about 120 segments. Color: ochre yellow in life (Hartman, 1941); light tan in alcohol. Prostomium entire on anterior margin; caruncle narrow, extending to end of setiger 2; no occipital tentacle; 2 pairs of eyes: anterior pair cup-shaped, posterior pair oval. Peristomium greatly inflated, extending anteriorly be- yond prostomium (Fig. 1A). Setiger 1 greatly reduced, but with capillaries in both noto- and neuro- podia (Fig. 1A). Setigers 2-4 with noto- and neuropodial fascicles of 468 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Y Vy Ge S YI WWE Z — — = WEE fy 7 —— (| Gf LEE CIEL TAL G Wh Ue A,G /}—— 500m ———_J B-D }—— 250um— E-F}— 3ym — Fig. 1. Carazziella citrona: A, Anterior end of holotype in dorsal view; B, Ventral major spine from setiger 5; C-D, Dorsal falcate major spines from setiger 5; E, Hooded hook from anterior setiger; F, Hooded hook from posterior setiger; G, Pygidium, in posterior view. unilimbate capillaries, neurosetae from setigers 6-9 unilimbate capillaries, these replaced by hooded hooks from setiger 10, hooks distally bifid in anterior setigers (Fig. 1E), secondary tooth gradually becoming smaller in middle and lost entirely in posterior setigers (Fig. 1F); hooks about 15-18 per fascicle in middle body region and accompanied throughout by superior and inferior capillaries. Setiger 5 distinctly modified, bearing modified spines and a ventral fascicle of unilimbate capillaries, dorsal fascicle of capillaries lacking. Major spines of 2 types: (1) ventral row with expanded bristled ends (Fig. 1B); (2) dorsal spines smaller, falcate, with bare tip emerging from dense cloak of bristles (Fig. 1C—D). Branchiae from setiger 7 (Fig. 1A), not overlapping at midline, continuing for over 70 setigers. Pygidium forming 4 subequal lobes (Fig. 1G). VOLUME 92, NUMBER 3 469 Remarks.—Carazziella citrona is unique among species of the genus in the nature of the posterior unidentate hooded hooks and that they begin on setiger 10. Carazziella hymenobranchiata Blake & Kudenov, 1978, from Victoria, Australia, bears hooded hooks from setiger 11-14, but in other species they first occur from setiger 7 or 8. Hartman (1941, pl. 45, fig. 13) described and figured the falcate spines of setiger 5 as being covered with both a hood and bristles. This appears to be an artifact of the angle of view and refraction of light. The curvature of the falcate spines often appears enlarged due to the crest of bristles; presumably Hartman mistook this for a hood. Hartman (1966) recorded Polydora citrona from Los Angeles Harbor. It seems possible, however, that this record may represent another species, possibly C. calafia; for this reason, it is not cited in the synonymy above, pending the relocation and examination of those specimens. Distribution.—California, Mission Bay (San Diego), intertidal in sandy mud areas inhabited by the thalassinid shrimp, Upogebia (Hartman, 1941). Carazziella hobsonae, new species Figs. 2-3 Material examined.—FLORIDA: Tampa Bay, 17 Oct. 1963, coll. John L. Taylor, dredged, 3 m.—holotype (USNM 56494) and 120+ paratypes (USNM 56495). TEXAS: 29°19'15"N; 94°38'42”W, 15 May 1975, coll. Texas A&M University, 13 m, sand-clay sediment.—3 paratypes (USNM 56496). Description.—A small species, up to 4.5 mm long and 0.5 mm wide for 40 setigerous segments. Color obscured due to staining by Rose Bengal; smaller individuals with irregular flecks of black pigment on dorsum, pre- sumably a vestige of larval pigment. Prostomium entire to vaguely incised on anterior margin (Fig. 2B); ca- runcle divided into 2 parts; anterior part extending posteriorly to middle of setiger 2, followed immediately by another discrete ridge abutting against anterior portion and extending to middle of setiger 3 (Fig. 2A—B); no occip- ital tentacle; 2 pairs of eyes: posterior pair closely spaced, oval; anterior pair widely spaced and cup-shaped. Peristomium well-developed, but not inflated; palps extend posteriorly to setiger 8. Setiger 1 with a dorsally situated fingerlike notopodial lobe lacking no- tosetae (Fig. 2A—B); neuropodium with well-developed fascicle of slender capillaries. Setigers 2, 3, 4, —, 6 and subsequent setigers with well-developed posteriorly directed fascicles of unilimbate capillary notosetae; capillaries of posterior notopodia longer, alimbate, no posterior spines. Neuropodia of setigers 2, 3, 4, -, 6 and 7 with short postsetal lobes and spreading fascicles of unilimbate capillaries; bidentate hooded hooks from setiger 8, accom- panied by capillaries throughout; anterior setigers with 3 hooks per ramus, 470 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON mM) it AN 3, equ —4 C-GH—— 250um—4 HL-— 10um_4 Fig. 2. Carazziella hobsonae: A, Anterior end in lateral view; B, Anterior end in dorsal view; C, Fascicle of major spines of setiger 5, in ventral view; D, Dorsal spines from setiger 5; E-F, Ventral spines from setiger 5; G, Superior dorsal capillary setae from setiger 5; H, Hooded hook; I, Pygidium in posterior view; J, Posterior end in dorsal view. gradually increasing to S—6 in middle, then decreasing to 2-3 in posterior setigers; hooks with only a slight angle between teeth (Fig. 2H). Setiger 5 distinctly modified, with a superior dorsal fascicle of 1 or 2 capillaries, 2 types of heavy bristled spines and a fully developed fascicle of unilimbate capillary neurosetae. Major spines arranged in 2 rows (Fig. 2C): (1) ventral row with 3—4 spines bearing expanded bristled tops, these appearing entire in ventral view (Fig. 2C), but with 2 knobs in lateral view, one distinctly smaller (Fig. 2E—F); (2) dorsal row with 2—4 smaller, falcate spines bearing bristles not reaching tip (Fig. 2D). Branchiae from setiger 7, continuing for 15—16 segments, these broad, straplike, overlapping at midline (Fig. 2B). Pygidium with 4 individual lobes (Fig. 2I-J). VOLUME 92, NUMBER 3 471 ESOPHAGUS INTESTINE | 100um GIZZARD Fig. 3. Carazziella hobsonae: A, Anterior part of digestive tract showing the structure and relationship of the gizzard to the esophagus and intestine. Digestive tract clearly visible through body wall, esophagus extending to about setiger 8, with a thickened muscular structure occupying setiger 9 (Fig. 3), followed by intestine [See also Blake, 1969, 1971, Blake & Wood- wick, 1972 and Blake & Kudenov, 1978 for a similar structure in several species of Polydora related to P. socialis and P. convexa; believed to func- tion as a gizzard]. | Remarks.—See comments for C. carrascoi. Ecology.—Carrazziella hobsonae occurs in soft sediments and is associ- ated with an echiuroid, Thallessena hartmanni and a bivalve, Paramya suborata off Texas (Mr. Clyde Henry, personal communication). Etymology.—This species is named in memory of the late Katharine D. Hobson, whose time with us was all too short, but whose excellent work on polychaetes will long be remembered. Distribution.—Florida; Texas. Carrazziella calafia, new species Fig. 4 Pseudopolydora reishi: Reish, 1968:84. [Not Woodwick, 1964] Material examined.—CALIFORNIA: Monterey Bay, coll. Moss Landing Marine Laboratories: 21 Aug. 1971, Sta. 1156, 36°53’N, 121°55’W, 37 m, grab No. 8.—1 paratype (CAS 00712); 21 Aug. 1971, Sta. 1157, 36°50.2'N; 121°50.2’W, 36.5 m, grab No. 2.—67 paratypes (USNM 56493); 21 Aug. 1971, Sta. 1158, 36°55’N; 121°5S6’W, 25.5 m, grab No. 5.—2 paratypes (CAS 00711): Elkhorn Slough, 15 June 1976, coll. J. A. Blake, approximately 250 m upstream from highway bridge, intertidal, sand-mud.—holotype (USNM 56491) and 1 paratype (CAS 00714); coll. Moss Landing Marine Laborato- ries: 20 July 1974, Sta. Sk 7, sand.—S paratypes (CAS 00713); Sta. Sk 6.— 1 paratype (USNM 56492); Los Angeles Harbor, coll. Harbors Environ- mental Projects: 21 May 1974, Sta. A2a, 10.7 m.—12 specimens (AHF); 20 472 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 4. Carazziella calafia: A, Anterior end in dorsal view; B—D, Ventral major spines from setiger 5; E-F, Dorsal major spines from setiger 5; G, Hooded hook; H, Posterior end in dorsal view. May 1974, Sta. A1l2, 11.3 m.—3 specimens (AHF); 21 May 1974, Sta. A9, 10.7 m.—8 specimens (AHF); 20 Aug. 1974, Sta. A2a, 10.7 m.—16 speci- mens (AHF); 7 May 1976, Sta. Alla, 6.1 m.—2 specimens (AHF); 5 Aug. 1976, Sta. A3a, 7.7 m.—2 specimens (AHF); | Dec. 1976, Sta. Cla, 12.8 m.—1 specimen (AHF); Santa Catalina Island, Indian Rock Reef, 12 m.— 5 specimens (AHF). MEXICO: Bahia de Los Angeles, coll. Beaudette Foundation: Oct. 1962, Sta. 85, silty sand.—3, specimens (USNM 57480); sta. 102, silty sand.—1 specimen (USNM 57481). Description.—A small species, up to 7 mm long and | mm wide for 48 setigerous segments. Color obscured on most specimens due to staining with Rose Bengal; no body pigment. Prostomium entire to weakly incised on anterior margin (Fig. 4A); carun- VOLUME 92, NUMBER 3 473 cle extending to anterior border of setiger 3; 4 eyes arranged as in C. hob- sonae. Peristomium well-developed, inflated; palps extending posteriorly for 7-9 setigers. Setiger 1 well-developed; prominent fascicles of capillaries in both rami of setigers 1-4; notopodia of setiger 6 and subsequent segments with fas- cicles of long capillaries; setigers 6-7 with capillary neurosetae; bidentate hooded hooks from setiger 8 (Fig. 4G) accompanied by unilimbate capillaries throughout; hooks numbering 4 per ramus in anterior setigers, gradually increasing to 6 in middle and posterior setigers. Setiger 5S distinctly modified, with a prominent fascicle of capillary neu- rosetae (Fig. 4A); no dorsal capillaries. Major spines of 2 types: (1) a ventral row with expanded tips bearing bristles and terminal depression (Fig. 4B— D); (2) dorsal row of falcate spines with bristles extending to end of spine and obscuring its tip (Fig. 4E—F). Branchiae broad, not meeting at midline, present from setiger 7 (Fig. 4A), continuing through middle body region. Pygidium with 4 lobes (Fig. 4H). Digestive tract with ‘‘gizzard’’ similar to that of C. hobsonae. Remarks.—See comments below under C. carrascoi. Ecology.—Intertidal to about 40 m, in substrata composed mostly of sand. Etymology.—The epithet originates from Calafia, the legendary black am- azon queen of an island utopia, as described by the sixteenth century Span- ish author Montalvo in Las Sergas de Esplandian, a continuation of his famous romance, Amadis de Gaula. California was the island where Calafia ruled. Distribution.—California: Monterey Bay, Los Angeles Harbor, Santa Catalina Island; Mexico: Bahia de Los Angeles (Gulf of California). Carazziella carrascoi, new species Fig. 5 Polydora citrona: Carrasco, 1974:193; 1976:25—28. [Not Hartman, 1941]. Material examined.—CHILE: Bahia de Concepcion, Jan. 1969, coll. V. A. Gallardo.—holotype (MZC 14652) and 2 paratypes (MZC 14653-4). Description.—A small species, up to 2.8 mm long and 0.4 mm wide for 25 segments for the type-material, but reported up to 7 mm long and 0.6 mm wide for 31 segments (Carrasco, 1974). Body light tan in alcohol with brown pigment along margin of caruncle. Prostomium flattened or rounded on anterior margin (Fig. 5A); caruncle extending to middle of setiger 2 thereupon terminating, followed immedi- ately by a discrete second ridge extending onto setiger 3; no occipital ten- tacle; 2 pairs of eyes: anterior pair cup-shaped, widely spaced; posterior pair oval, closely spaced. Peristomium inflated, extending anteriorly to and, 474 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 5S. Carazziella carrascoi: A, Anterior end in dorsal view; b, Hooded hook; C-D, Ven- tral major spines from setiger 5; E-F, Dorsal major spines from setiger 5; G, Posterior end in lateral view; H, Pygidium in posterior view. sometimes, beyond prostomium; palps extending posteriorly for 7-8 setig- ers. Setiger 1 with notopodial lobes significantly longer than neuropodial lobes; with 2-3 very thin, short capillary notosetae and well-developed fascicle of prominent unilimbate neuropodial capillaries. Setigers 2-4 with fascicles of unilimbate capillaries in both rami, from setiger 6, long, thin, unilimbate capillary notosetae; neuropodia of setigers 6 and 7 bearing cap- VOLUME 92, NUMBER 3 475 illaries only; bidentate hooded hooks first appearing in neuropodia from setiger 8 (rarely 9) (Fig. 5B), these numbering 3-5 per fascicle, accompanied by 2-3 inferior capillaries and occasionally, also by a single superior cap- illary, for most of body length. Setiger 5 distinctly modified, with heavy dorsal musculature overlapping onto setiger 6; with or without a simple superior dorsal seta, and bearing a prominent fascicle of bilimbate neurosetae. Major spines of 2 types in a double curved row: (1) ventral row of large spines distally expanded into 2 indistinct rounded knobs, one more distal to the other in lateral view. (Fig. 5C-D); (2) dorsal row of smaller, falcate spines with bristles surrounding shaft subdistally, leaving tip free (Fig. 5E—F). Branchiae from setiger 7, not overlapping at midline, continuing for 6-8 setigers (Fig. SA). Pygidium with 4 conical lobes, the dorsal pair less than half as large as the ventral pair (Fig. 5G—H). A highly muscular gizzard present in gut at level of setigers 7-9. Remarks.—Carazziella hobsonae, C. calafia and C. carrascoi are closely related and form a distinct species-group, characterized by having a ‘‘giz- zard’’ in the digestive tract. This structure has not been observed in the other 8 species of the genus. The caruncle is divided in C. hobsonae and C. carrascoi, and undivided in C. calafia. Notosetae are present on setiger 1 of both C. calafia and C. carrascoi: the former species has a well-devel- oped fascicle, whereas the latter has only 2—3 very delicate setae. C. hob- sonae lacks notosetae on setiger 1 altogether. The superior dorsal fascicle is absent from setiger 5 in C. calafia, present in C. hobsonae, and may be present or absent in C. carrascoi. Each species differs from the others in the configuration of the major spines of setiger 5 (compare Figs. 2C—F; 4B-— F; 5C—F). Carazziella hobsonae and C. calafia each have a terminal depres- sion on these setae; this is lacking on C. carrascoi. Etymology.—This species is named in honor of Dr. Franklin D. Carrasco, polychaete systematist of the Instituto de Biologia (MZC), who supplied the specimens on which this description is based. Distribution.—Chile, Bahia de Concepcion, subtidal in mixed sediments. Carazziella patagonica, new species Fig. 6 Material examined.—ARGENTINA: Golfo San Matias, Sta. SAO I-49, 41°16’S; 65°04’ W, coll. 1971, J. Orensanz, 36 m, mixed sand-mud substra- tum.—holotype (MACN 29262). Description.—Holotype small, measuring 1.7 mm long and 0.27 mm wide for 25 setigers. Color light tan, with a few small black medial pigment spots. Prostomium entire on anterior margin; with caruncle extending poste- riorly into setiger 2; no occipital tentacle. but posterior part of caruncle 476 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON \ N\ Vy ih Na A,H #—100um———J B-G -—10um—4 Fig. 6. Carazziella patagonica: A, Anterior end in dorsal view; B, Hooded hook; C-D, Dorsal major spines from setiger 5; E-G, Ventral major spines from setiger 5; H, Posterior end in dorsolateral view. forming a fold overlapping anterior portion of caruncle (Fig. 6A); 2 pairs of eyes arranged in wide crescent; lateral pair larger, cup-shaped; medial pair smaller, rounded. Peristomium greatly enlarged; palps missing. Setiger | reduced, lacking distinct segmental boundaries; neuropodium completely absent; notopodium with short postsetal lobe bearing 2-3 thin capillaries (Fig. 6A). Setigers 2, 3, 4, —, 6 and subsequent segments with well-developed notopodia bearing unilimbate capillaries with finely granu- lated shafts and thin transparent sheaths. Similar neurosetae in setigers 2, 3, 4, -, 6 and 7; bidentate hooded hooks in neuropodia from setiger 8 (Fig. 6B) numbering 3—4 per ramus, accompanied by 2 thin inferior capillaries. VOLUME 92, NUMBER 3 477 Setiger 5 distinctly modified, bearing a single unilimbate, granulated su- perior dorsal capillary and a ventral fascicle of 4—5 unilimbate nongranulated capillaries. Major spines including 2 dorsal falcate spines bearing a distinct crest of bristles partly surrounding the tip (Fig. 6C—D) and 3-4 ventral api- cally expanded spines, each surmounted by a tall cone of bristles (Fig. 6E-G). Branchiae from setigers 7-10, each gill long, straplike (Fig. 6A), gills meeting middorsally. Pygidium with 2 small cirri (Fig. 6H), but specimen may be damaged. Remarks.—The reduction of setiger 1 in C. patagonica is unique in the genus. No other species in the entire Polydora—complex has a well-developed notopodium with setae on setiger 1 while entirely lacking a neuropodium. Etymology.—The epithet is derived from Patagonia, the general regional name applied to the southern half of Argentina. Distribution.—Argentina, in shallow subtidal sediments. Carazziella reishi (Woodwick, 1964) Fig. 7 Pseudopolydora reishi Woodwick, 1964:152; Kohn & Lloyd, 1973:381. [Not Reish, 1968:84]. Material examined.—ENIWETOK: Engebi, 7 Sept. 1956, coll. D. J. Reish, hard coral rocks, in sand.—holotype (USNM 32611). INDONESIA: Pulo Boonda, NW of Sumatra, 5°33’N; 95°09’E, TeVega Sta. 93, Nov. 1963, coll. A. J. Kohn, coral rocks.—4 specimens (USNM 45318). JOHNSTON ATOLL: Sta. 3-5, 25 April 1976, coll. R. Brock.—98 specimens (NOSC 77003051). Description.—A small species, up to 3.0 mm long and 0.2 mm wide for 35 setigers. Body light tan in alcohol with no pigment. Prostomium incised on anterior margin (Fig. 7A); caruncle extending to end of setiger 2; no occipital tentacle; no eyes. Peristomium well-developed, but not inflated; palps missing from all material examined. Setiger 1 well developed, with conical postsetal lobes in both noto- and neuropodia, each lobe with fascicle of capillaries (Fig. 7A); setigers 2, 3, 4, —, 6 and subsequent setigers with well-developed fascicles of unilimbate capillary notosetae; no specialized posterior spines. Neuropodia of setigers 2, 3, 4, -, 6 and 7 with short conical postsetal lobes and spreading fascicles of unilimbate capillaries; 2—3 bidentate hooded hooks beginning on setiger 8, accompanied throughout by 1-2 inferior capillaries; margin of hood ser- rated (Fig. 7B). Setiger 5 distinctly modified, with a superior dorsal fascicle of 2—3 cap- illaries located anterodorsal to the major spines; with a prominent fascicle of unilimbate neurosetal capillaries. Major spines arranged in 2 curved rows: 478 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON ( Mi Wy Y AN i hy i we, Spe G PANO Ww SS ay —<—$— aC 5 7 ane Fig. 7. Carazziella reishi: A, Anterior end in dorsal view; B, Hooded hook; C-D, Ventral major spines from setiger 5; E, Falcate dorsal spine from setiger 5; F, Pygidium in posterior view. (1) a ventral row of 3 spines with expanded bristled tops bearing a small barely visible apical cone (Fig. 7C—D); (2) a dorsal row of 2—3 simple falcate spines, lacking bristles (Fig. 7E). Branchiae from setiger 7, continuing for only 5 setigers; each gill broad, reaching to the midline, with 6 patches of cilia extending in a line across dorsum between each gill pair (Fig. 7A). Pygidium with 4 short thickened lobes (Fig. 7F), these not conical as in most species of genus. VOLUME 92, NUMBER 3 479 Remarks.—This species was well described by Woodwick (1964). How- ever, the new material from Indonesia (see Kohn & Lloyd, 1973) and John- ston Atoll has permitted a more detailed study of the morphology of this form for comparison with the now numerous other species of the genus. As a result of these studies, it was noted that the hooded hooks on the Indo- nesian and Johnston Atoll specimens bear fine serrations on their hood Openings; such serrations were not observed on the holotype due to the small size of that specimen and its setae, precluding observation of that character with light microscopy. The bristle-topped spines of setiger 5 have a small central cone among the bristles of all specimens examined. The 6 ciliary patches located on the dorsum between each gill pair were clearly visible on the Indonesian and Johnston Atoll specimens, but only partially discernible on the holotype because it was dorsally damaged. The pygidium is definitely composed of 4 lobes, and not 2 as stated by Woodwick (1964). Carazziella reishi is unique within the genus in exhibiting enlarged bristle- topped spines accompanied by unbristled falcate spines. In 9 other species (Blake & Kudenov, 1978; this paper) both types of spines are bristle-topped. In C. quadricirrata (Rainer, 1973) both the cusped spines and the simple, falcate spines lack bristles. C. reishi has less of an oblique angle between the main fang and shaft of the hooded hooks than other species of the genus, is the only species known to have serrations on the opening of the hooded hooks, discrete ciliary patches between the gill pairs and is the only species lacking eyes. Moreover, C. reishi is the only species of the genus reported to bore into calcareous substrata. Distribution.—Indonesia; Johnston Atoll; Enewetak. DISCUSSION The Polydora-complex has been revised by Blake & Kudenov (1978) to include 6 genera: Polydora Bosc. 1802; Pseudopolydora Czerniavsky, 1881; Carazziella Blake & Kudenov, 1978; Tripolydora Woodwick, 1964; Boc- cardiella Blake & Kudenov, 1978 and Boccardia Carazzi, 1893. The first 3 genera have branchiae beginning posterior to setiger 5, while the last 3 bear branchiae beginning on setiger 2. Blake & Kudenov (1978) present a com- plete review of the characters which separate and distinguish these 6 genera. Woodwick (1964) was the first to observe that Polydora citrona Hartman is a species which does not completely agree with the generic definition of Polydora. Blake & Kudenov (1978) designated P. citrona as the type- species of Carazziella which now contains the following 11 species, listed geographically: C. victoriensis Blake & Kudenov, 1978. Victoria, Australia C. phillipensis Blake & Kudenov, 1978. Victoria, Australia C. hymenobranchiata Blake & Kudenov, 1978. Victoria, Australia 480 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON . hirsutiseta Blake & Kudenov, 1978. New South Wales, Australia quadricirrata (Rainer, 1973). New Zealand . reishi (Woodwick, 1964). Enewetak; Indonesia; Johnston Atoll citrona (Hartman, 1941). California calafia, new species. California; western Mexico . hobsonae, new species. Gulf of Mexico . carrascoi, new species. Chile . patagonica, new species. Argentina. aaaae eee Only 2 of these species, C. hobsonae and C. patagonica, occur outside of the Pacific basin. Of the 11 species, C. hymenobranchiata, C. hirsutiseta, C. quadricirrata, C. citrona, C. reishi and C. patagonica are specifically unique and readily separable from one another. Carazziella victoriensis, C. phillipensis, C. hobsonae, C. calafia and C. carrascoi, however, are all closely related. Carazziella victoriensis and C. phillipensis from Australia, lack a gizzard between the esophagus and intestine, whereas C. hobsonae, C. calafia and C. carrascoi, all from American waters, possess such a gizzard. Species within these 2 groups may be distinguished from one another by considering the presence or absence of notosetae on setiger 1, shape of the prostomium and caruncle, presence or absence of superior dorsal capillaries on setiger 5 and the form of the major spines of setiger 5. Each of these species is small and careful attention to detail is required for accurate identification. The occurrence of a gizzard in the C. hobsonae group Is interesting. The structure was first reported from larvae and adults of Polydora socialis (Blake, 1969; 1971). Blake & Woodwick (1972) report a similar structure in P. convexa. Blake & Kudenov (1978) report the occurrence of gizzards in P. socialis, P. protuberata and P. tentaculata, all of which are very closely related. The structure also occurs in the closely related P. cardalia (Blake, in preparation). In each genus, whether Polydora or Carazziella, the struc- ture consists of 4 symmetrically arranged muscles which apparently serve to crush or otherwise treat food materials as they pass from the esophagus to the intestine. The specimens of C. hobsonae permit a clear look at the gross morphology of this structure (Fig. 3) and there appears to be a valve which may control entry of materials into the gizzard. A histological study is needed to assess the fine structure and function of this gizzard. The gizzard is easily overlooked in routine investigation, especially when search- ing for the other more obvious external taxonomic characters. Literature Cited Blake, J. A. 1969. Reproduction and larval development of Polydora from northern New England (Polychaeta: Spionidae).—Ophelia 7: 1-63. —. 1971. Revision of the genus Polydora from the east coast of North America (Poly- chaeta: Spionidae).—Smithsonian Contrib. Zool. 75: 1-32. VOLUME 92, NUMBER 3 481 , and J. D. Kudenov. 1978. The Spionidae (Polychaeta) from southeastern Australia and adjacent areas with a revision of the genera.—Mem. Nat. Mus. Victoria 39:171- 280. , and K. H. Woodwick. 1972. New species of Polydora (Polychaeta: Spionidae) from the coast of California.—Bull. South. California Acad. Sci. 70:72-79. Carrasco, F. D. 1974. Spionidae (Polychaeta) provenientes de la Bahia de Concepcion y lugares adyacentes.—Bol. Soc. Biol. Concepcion 48:185-—201. 1976. Larvas de la familia Spionidae (Polychaeta) en el plancton de la Bahia de Concepcion, Chile.-—Gayana (Zoologia), No. 38:1-63. Hartman, O. 1941. Some contributions to the biology and life history of Spionidae from California.—Allan Hancock Pacific Expeds. 7(4):289-324, pls. 45-48. . 1961. Polychaetous annelids from California.—Allan Hancock Pacific Expeds. 25:1- 226, 34 pls. . 1966. Quantitative survey of the benthos of San Pedro Basin, southern California. Pt. 2. Final results and conclusions.—Allan Hancock Pacific Expeds. 19(2): 187-456, 14 pls., 2 maps. . 1969. Atlas of the sedentariate polychaetous annelids from California——Allan Han- cock Foundation, University of Southern California, Los Angeles, 812 pp. Kohn, A. J., and M. C. Lloyd. 1973. Polychaetes of truncated reef limestone substrates on eastern Indian Ocean coral reefs: diversity, abundance and taxonomy.—Int. Rev. Ges. Hydrobiol. 58:369—400. Rainer, S. 1973. Polydora and related genera (Polychaeta: Spionidae) from Otago waters.— Jour. Roy. Soc. New Zealand 3:545-564. Reish, D. J. 1968. A biological survey of Bahia de Los Angeles, Gulf of California, Mexico. II. Benthic polychaetous annelids.—Trans. San Diego Soc. Nat. Hist. 15(7):67—106. Woodwick, K. H. 1964. Polydora and related genera (Annelida, Polychaeta) from Eniwetok, Majuro and Bikini Atolls, Marshall Islands.—Pacific Science 18:146—159. Tomales Bay Marine Laboratory, Marshall, California 94940. PROC. BIOL. SOC. WASH. 92(3), 1979, pp. 482-493 THE INDO-PACIFIC PIPEFISH GENERA NOTIOCAMPUS GEN. NOV. AND NANNOCAMPUS GUNTHER C. E. Dawson _Abstract.—Notiocampus (type-species: Nannocampus ruber Ramsay and Ogilby) is described from Australia. A revised diagnosis is given for the superficially similar Nannocampus Gunther, and its recognized species (N. subosseus Gunther, N. elegans Smith) are redescribed; all treated species are illustrated. Study of an Australian pipefish identified as Nannocampus ruber Ramsay and Ogilby showed it to differ markedly from other species of Nannocampus Gunther, and this prompted examination of other material referred to this poorly known syngnathine (tail pouch) genus. As a result, I here redescribe N. ruber, refer it to a new genus, and provide a revised diagnosis of Nan- nocampus together with brief descriptions of the two species therein rec- ognized. Material is, in most cases, limited, and additional specimens may eventually warrant emendation of diagnoses and descriptions. However, present data clarify or correct early descriptions and should facilitate future study of these relatively uncommon Indo-Pacific pipefishes. Taxa treated here are in part characterized by the lack of pectoral and anal fins in sub- | adults and adults, but pectoral fins may well be present in larvae. Dawson and Allen (1978) have shown that certain pipefishes that lack dorsal and pectoral fins as adults have these fins developed in pouch and/or planktonic larvae. Similarly, pectoral fins are present in planktonic larvae but absent in adults of the North Atlantic genus Nerophis. In the absence of larval specimens, present generic diagnoses apply only to juveniles or adults. Counts and measurements follow Dawson (1977) and are based on the material examined; color descriptions are from specimens preserved in al- cohol. Abbreviations for repositories of examined material follow: AMS, Australian Museum, Sydney; BMNH, British Museum (Natural History); GCRL, Gulf Coast Research Laboratory Museum; NMW, Naturhisto- risches Museum, Wien; SAM, South Australian Museum, Adelaide; WAM, Western Australian Museum, Perth; ZMA, Zoologisch Museum, Amster- dam. Notiocampus, new genus Type-species.—Nannocampus ruber Ramsay and Ogilby, 1886. Diagnosis.—Superior trunk and tail ridges confluent; lateral tail ridge ab- VOLUME 92, NUMBER 3 483 sent; inferior trunk and tail ridges discontinuous near anal ring; lateral trunk ridge confluent with inferior tail ridge; body slender, elongate, its depth 3- 4 in head length (HL); trunk nearly round in section; venter but slightly V-shaped, without prominent median ridge or keel; dorsum and venter of tail somewhat convex; scutella inconspicuous, without longitudinal keels, their width less than half of ring length. Head length 15—16 in standard length (SL); median dorsal snout ridge low, entire, originates on posterior half of snout, terminates on interorbital; opercle smooth to minutely striate; head elsewhere without prominent ridges or ornamentation; body ridges and rings inconspicuous, devoid of spines, serrations or dermal flaps; dorsal-fin origin on tail, fin base not elevated, its length more than 3 in HL; pectoral and anal fins absent, caudal fin present; total rings 66-68; nares with two widely spaced pores of unequal diameter; brood pouch unknown, presumably sub- caudal; without odontoid processes in jaws (Dawson and Fritzche, 1975). Maximum size at least 151 mm SL; Indo-Pacific, marine. Comparisons.—Among pipefishes with confluent superior trunk and tail ridges, Notiocampus exhibits a unique combination of characters (e.g. pec- toral and anal fins absent, caudal fin present, dorsal fin located on tail, confluent lateral trunk and inferior tail ridges). This ridge configuration and absence of pectoral and anal fins is shared with adults of Penetopteryx Lunel and Apterygocampus Weber but those forms lack the dorsal fin in adults and have total ring counts of 45-61 (dorsal fin present in Notiocampus, total rings 66-68). The North Atlantic or European Entelurus Duméril and Ne- rophis Rafinesque also lack pectoral and anal fins in adults but Entelurus has 28-31 trunk rings and the dorsal fin is mainly on the trunk (trunk rings 18-19, dorsal fin on tail in Notiocampus). Further, Nerophis lacks the cau- dal fin (present in Notiocampus). Nannocampus shares the fin complement of Notiocampus but total rings are less (47-56 against 66-68) and inferior trunk and tail ridges are confluent rather than discontinuous. Etymology.—From the Greek notios (southern) and kampos (sea-animal), in allusion to the southern hemisphere habitat of the type-species. Gender: masculine. Remarks.—To my knowledge, there are now only three specimens refer- able to this genus in collections. It is evident that these forms are not com- monly taken by usually employed methods (nets, ichthyocide) and they may prove to be cryptic species wherein adults occupy the interstices of rock or coral rubble. Notiocampus ruber (Ramsay and Ogilby) Figs. 1 and 2 Nannocampus ruber Ramsay and Ogilby, 1886:757 (original description; Shark Reef, Port Jackson, New South Wales, Australia). Diagnosis.—See generic diagnosis. 484 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON RISES Terao a VOLUME 92, NUMBER 3 485 Fig. 2. Notiocampus ruber. Top: AMS B.9199 (110 mm SL, holotype). Bottom: AMS 1.20167-013 (151 mm SL). Description.—Counts and measurements (mm) of 110 mm SL holotype are followed (in parentheses) by those of 151 mm SL specimen from Kan- garoo I.: trunk rings 19 (18), tail rings 49 (48), dorsal-fin rays 11 (13), total subdorsal rings 1.5 (1.25), caudal-fin rays 6 (7), HL 7.5 (9.3), snout length 2.9 (3.3), snout depth 0.9 (1.0), length of dorsal-fin base 2.1 (2.6), depth at anal ring 1.9 (2.7), trunk depth 2.0 (2.9). Dorsal-fin origin at rear margin of Ist tail ring in holotype, about '4 of ring length behind anterior margin of Ist tail ring in 151 mm fish and at anterior margin of Ist tail ring in a 3rd (damaged) specimen. Median dorsal snout ridge terminates on interorbital, <_— Fig. 1. Notiocampus ruber. Upper pair—Lateral and dorsal aspects of head and anterior trunk rings with detail of left naris. Lower pair—Top: Posterior trunk and anterior tail rings illustrating ridge pattern and dorsal fin; arrow locates anal ring. Bottom: Posterior tail rings and caudal fin with diagrams of selected section areas. From AMS 1.20167-013. 486 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON not confluent with orbital ridges. Nares (Fig. 1), bilaterally, with large simple pore above, well separated from a minute pore on slender anteroventral tubiform process. Posterior tail rings not narrowed above and below to form keel-like ridge. The fresh holotype was described as ‘‘red with some minute white spots.”’ The holotype and Kangaroo I. specimen are now light tan without evidence of spots or other persistent markings. Remarks.—The three study specimens are presumably females. Whitley (1941) reported the presence of ‘‘thick mucus canals’’ on the head of the holotype. I find no such canals, nor is there evidence of a mucous coating which often occurs in some preserved pipefishes, especially in species of Corythoichthys Kaup. The holotype, now in two pieces, has the dorsal fin adnate to dorsum and fin rays are difficult to count with certainty. There seem to be 11 rays as originally described but 12 pterygiophores may be present. The specimen reported by Glover (1968) is now dried, part of the tail is missing, and dorsal-fin rays cannot be counted accurately. Although noting differences in dorsal-fin length and a “‘reduced’’ caudal fin, Whitley (1941) mistakeniy referred a specimen labeled ‘‘South Head, Sydney’’ to Nannocampus ruber. This fish (AMS IB.560), now 129 mm TL, has 19 + 54 rings, 29-30 rays in the damaged dorsal fin and 2 + 5.25 sub- dorsal rings; median dorsal snout ridge confluent with supraorbital ridges; superior body ridges confluent; lateral trunk ridge indistinct, possibly con- fluent with inferior tail ridge; pectoral and anal fins absent; caudal fin absent; last 5—6 tail rings poorly defined, gradually decreasing in length, laterally compressed and margined above and below by a keel-like ridge which con- tinues around tip of tail. This specimen clearly differs from N. ruber in a number of characters, whereas general morphology and counts agree closely with Nerophis lumbriciformis (Jenyns). This fish may represent an unde- scribed Australian species, but I believe that it is a mislabeled Nerophis of European origin. Comparisons.—See generic diagnosis. Distribution.—Known only from New South Wales and South Australia. One specimen (SAM F.3220) was taken in a “‘craypot,’’ another (AMS [.20167-013) taken with SCUBA from a ‘‘rocky ledge’’ at a depth of 15 meters. Material examined.—AMS B.9199 (110 mm SL, presumably female, ho- = Fig. 3. Nannocampus subosseus. Upper pair—Lateral and dorsal aspects of head and an- terior trunk rings. Lower pair—TJop: Posterior trunk and anterior tail rings illustrating ridge pattern and dorsal fin. Bottom: Posterior tail rings and caudal fin. From WAM P.25758-023 (77.5 mm SL, female). 487 itiven VOLUME 92, NUMBER 3 488 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON lotype), Australia, New South Wales, Port Jackson, Shark Reef, 1885. AMS 1.20167-013 (1451 mm SL), South Australia, Kangaroo I., 35°37’S, 137°15’E, 1978, H. Larson. SAM P.3220 (ca. 130 mm, damaged), South Australia, Cape Jaffa, 1964, I. R. McInnes. Nannocampus Gunther Nannocampus Gunther, 1870:178 (type-species by original designation: Nannocampus subosseus Gunther, 1870). Diagnosis.—Superior trunk and tail ridges confluent; lateral tail ridge ab- sent; inferior trunk and tail ridges confluent; lateral trunk ridge ends, with or without deflection ventrad, near anal ring; body moderately slender, its depth 2.4—3.2 in HL; trunk not round in section, venter V-shaped, without median ridge or keel; dorsum of tail mainly flat, venter slightly convex; scutella inconspicuous, without longitudinal keels, their width less than half of ring length. Head length 10-15 in SL; median dorsal snout ridge low to distinctly elevated above dorsal margin of orbit, entire, originates on ante- rior third of snout, confluent behind with anterior continuations of supraor- bital ridges; opercle smooth to finely striate or pocked with minute depres- sions; head elsewhere without prominent ridges or ornamentation; body ridges and rings inconspicuous, devoid of spines, serrations or dermal flaps; dorsal-fin origin near anal ring, fin base somewhat depressed between su- perior ridges; pectoral and anal fins absent; caudal fin rather well developed, usually with 10 fin rays; total rings 47-56; nares with 2 separated but not widely spaced pores of subequal diameters; brood pouch subcaudal, pouch plates and dermal folds present, pouch closure unknown; without odontoid processes in jaws. Maximum size at least 110 mm SL; Indo-Pacific, marine. Comparisons.—See under Notiocampus above. Nannocampus subosseus Gunther Figs. 3 and 4 Nannocampus subosseus Gunther, 1870:178 [original description; Freyci- net’s Harbour (Western Australia)]. Nannocampus weberi Duncker, 1915:99 [original description; Malé Kuba Bay, Sumba I. (Indonesia)]. Diagnosis.—Lateral trunk ridge not deflected near anal ring; total rings 47-50; dorsal-fin rays 14-18. Description.—Rings 15 + 32-35; total subdorsal rings 3.0—4.25; dorsal-fin origin on posterior '4 of anal ring in holotype, at rear margin of anal ring and anterior 34 of Ist tail ring in two other Australian specimens. Median dorsal snout ridge usually elevated above dorsal rim of orbit; lateral trunk ridge ends between middle and posterior margin of anal ring. VOLUME 92, NUMBER 3 ee DS Age7 geo ee Meotl Lb4 277 Fig. 4. Upper pair: Nannocampus subosseus WAM P.25758-023 (77.5 mm SL, female). Lower pair: Nannocampus elegans GCRL 15963 (top, 101 mm SL, male; bottom, 93.5 mm SL, female). 490 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Ground color brown to dark brown; lower jaw and maxilla brown; pre- maxilla, median snout ridge and most of dorsum of head pale to tan; side and venter of snout crossed by broad near-white band extending below level of nares; upper half of orbit margined by series of short dark brown bars, of which one crosses interorbital; upper 4 of opercle brown with irregular short dark brown streaks, remainder of opercle and underside of head brown; trunk and tail with about six diffuse dark margined brown bars crossing dorsum and sides, bars about 2 of ring width and spaced 5-6 rings apart; upper '4 of side and dorsum pale between 13th and 20th rings; pos- terior 4% of tail mainly light tan with irregular darker bars between rings; dorsal-fin rays flecked with brown; caudal fin brown with pale margin (color description from WAM P.25758-023). Comparisons.—See this section under N. elegans. Remarks.—Gunther (1870), Duncker (1915) and Weber and de Beaufort (1922) failed to describe the lateral trunk ridge of the dried holotype (BMNH 1868.12.27.57), but Palmer (1954) reported that the specimen had been trans- ferred to alcohol and that the lateral ridge ends on the last trunk ring. The holotype, now 85 mm (originally 90 mm total length), lacks some posterior tail rings and the caudal fin, retains no evidence of original coloration, and is preserved in alcohol. Well-developed pouch plates extend below 13 tail rings but pouch folds have evidently been lost during preservation. My counts and measurements (mm) follow: trunk rings 15, tail rings remaining 33, subdorsal rings 0.25 + 2.75, HL 7.4, snout length 2.0, snout depth 2.0, length of dorsal-fin base 4.8, anal ring depth 3.1, trunk depth 3.4. Weber (1913) reported a specimen (as subosseus) from Indonesia which Duncker (1915) described (without examination) as N. weberi on the basis of differences from subosseus in published counts of rings and subdorsal rings. My counts and measurements (mm) of the 54.5 mm SL holotype of N. weberi follow: rings 15 + 32, dorsal-fin rays 18, subdorsal rings 0.5 + 3.75, caudal-fin rays 10, HL 5.4, snout length 1.6, snout depth 1.1, length of dorsal-fin base 4.4, anal ring depth 2.1, trunk depth 2.7. This presumably juvenile specimen generally agrees with examined N. subosseus in all char- acters except the tail ring count (32 against 35 in undamaged subosseus) and in the lateral configuration of the snout. The dorsal margin of the snout is straight to somewhat convex in N. subosseus (ca. 71-90 mm SL), whereas the margin is convex in the holotype of N. weberi and the snout ridge fails to reach the level of the dorsal rim of the orbit (reaches to or above rim in other subosseus). The difference in the tail ring count is within expected variation and the concave snout may reflect the juvenile condition. In the absence of additional distinguishing features and lack of adequate study material, I provisionally refer N. weberi to the synonymy of N. subosseus. Distribution.—Western Australia and Indonesia. One Australian speci- men was taken in a “‘rockpool’’; the Indonesian fish was collected ‘‘on reef.”’ 491 VOLUME 92, NUMBER 3 4 feee4 . Upper pair—Lateral and dorsal aspects of head and anterior Nannocampus elegans . Lower p Fig. 5. ridge pattern and dorsal ing d caudal fin. From GCRL 15965 (82 mm SL, female). rings illustrati l k and tai rior trun Poste . i air—Top rings trunk fin. Bottom rings an ] rior tal Poste 492 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Material examined.—Western Australia: BMNH 1868.12.27.57 (85 mm, damaged male, holotype), Freycinet’s Harbour, HERALD; WAM P.25758- 023 (1, 77.5), Rottnest I., Fish-hook Bay, 8 Mar. 1977, B. Hutchins; WAM P.25842-001 (1, 71), Houtman Abrolhos, Beacon I., rockpool 29 Aug. 1977, A. Lovell and H. Merrifield. Indonesia: ZMA 104.659 (54.5, juvenile, ho- lotype of N. weberi), Sumba (Soemba) I., Bay of Malé Kava, on reef, 10 Feb. 1909, van der Sande. Nannocampus elegans Smith Figs. 4 and 5 Nannocampus (sp.) nov. Smith, 1951:53 (notice of undescribed species; Xora River to Inhambane, South-east Africa). Nannocampus elegans Smith, 1961:516, fig. 355a [original description; Xora mouth northwards (SE Africa)]. Diagnosis.—Lateral trunk ridge deflected ventrad near anal ring; total rings 54-56; dorsal-fin rays 21-22. Description.—Rings 14 + 40-42, subdorsal rings 2.0—1.25 + 3.75— 4.25 = 5.5-6.0, caudal-fin rays 10 (see Smith, 1963 for additional counts). Proportional data based on five specimens, 56—106.5 mm SL (GCRL 15963-— 65), follow: HL in SL 12.4—-15.0, snout length in HL 3.0-3.6, snout depth in snout length 1.1—1.8, length of dorsal-fin base in HL 0.7-0.9, anal ring depth in HL 2.4—3.2, trunk depth in HL 1.7—2.4. Median dorsal snout ridge low, not elevated to or above dorsal rim of orbit, snout concave in lateral profile; lateral trunk ridge deflected variably ventrad, may reach but not confluent with inferior ridge. Ground color tan to brown; plain, or head and sides and dorsum of body irregularly streaked with fine brown lines or blotches; dorsal and caudal rays flecked with brown. For color in life see Smith, 1963. Comparisons.—Higher counts of total rings (54-56 against 47-50) and dorsal-fin rays (21-22 against 14-18) as well as a more anterior dorsal-fin insertion (on 2.0—1.25 trunk rings against 0.5 trunk—0.75 tail rings) distin- guish N. elegans from N. subosseus. The lateral trunk ridge is deflected to a greater or lesser degree in all elegans examined whereas this ridge ends without deflection in subosseus. In addition, the snout ridge is consistently low in elegans (distinctly elevated in subosseus) and preserved specimens of elegans lack the persistent pattern of bars and pale blotches found in subosseus. Distribution.—According to Smith (1963), this species is not uncommon in tidepools from South Africa (34°S) to Inhaca, Mozambique. Material examined.—South Africa: GCRL 15963 (2, 93.5-101); GCRL 15964 (1, 56); GCRL 15965 (2, 82—106.5); NMW 75533 (2, 78—96.5). VOLUME 92, NUMBER 3 493 Acknowledgments For permission to examine types or other specimens in their care I thank G. R. Allen and B. Hutchins (WAM), C. J. M. Glover (SAM), R. Hacker and P. Kahsbauer (NMW), H. Nijssen (ZMA), J. R. Paxton and D. F. Hoese (AMS) and A. C. Wheeler (BMNH). Special acknowledgment is due Mar- garet Smith, J. L. B. Smith Inst. of Ichthyology, Rhodes Univ., for the donation of specimens of N. elegans. Drawings are by Nancy Gordon. This study was supported in part by National Science Foundation Grant BMS 75-19502. Literature Cited Dawson, C. E. 1977. Review of the pipefish genus Corythoichthys with description of three new species.—Copeia [977:295-338. , and G. R. Allen. 1978. Synopsis of the “‘finless’’ pipefish genera (Penetopteryx, Ap- terygocampus and Enchelyocampus, gen. nov.).—Rec. West. Austr. Mus. 6(4):391—411. , and R. A. Fritzsche. 1975. Odontoid processes in pipefish jaws.—Nature 257:390. Duncker, G. 1915. Revision der Syngnathidae.—Mitteil. Naturh. Mus. Hamburg 32:9-120. Glover, C. J. M. 1968. Further additions to the fish fauna of South Australia.—Rec. S. Austr. Mus. 15(4):791-794.. Gunther, A. 1870. Catalogue of the fishes in the British Museum. Vol. 8. 549 pp. London. Palmer, G. 1954. Notes ona collection of syngnathid fishes from Malaya.—Bull. Raffles Mus. 25:27-30. : Ramsay, E. P., and J. Douglas-Ogilby. 1886. Description of two new fishes from Port Jack- son.—Proc. Linn. Soc. New South Wales 10:757-758. Smith, J. L. B. 1951. Thirteen noteworthy additions to the south-east African marine fauna.— Ann. Mag. Nat. Hist.,. ser. 12, 4(37):49-66. . 1961. The sea fishes of southern Africa. 4th edn.—Central News Agency Ltd., South Africa. —. 1963. Fishes of the family Syngnathidae from the Red Sea and western Indian Ocean.—Ichthyol. Bull. Rhodes Univ. 27:515-543. Weber, M. 1913. Die Fische der Siboga-Expedition.—Vol. 57 in Siboga Expedition Report, Leiden. 710 pp. , and L. F. de Beaufort. 1922. The fishes of the Indo-Australian Archipelago. Vol. 4. 410 pp.—E. J. Brill, Leiden. Whitley, G. P. 1941. Ichthyological notes and illustrations.—Austr. Zool. 10(1):1—50. Gulf Coast Research Laboratory Museum, Ocean Springs, Mississippi 39564. PROC. BIOL. SOC. WASH. 92(3), 1979, pp. 494-497 AN OLD WORLD OCCURRENCE OF THE EOCENE AVIAN FAMILY PRIMOBUCCONIDAE Storrs L. Olson and Alan Feduccia Abstract.—Parvicuculus minor Harrison and Walker, from the Lower Eocene of England was incorrectly assigned to the Cuculidae (Cuculiformes) by its authors and is here referred to the Primobucconidae, a family hitherto known only from the Lower and Middle Eocene of Wyoming. Procuculus minutus Harrison and Walker, also from the Lower Eocene of England, belongs neither in the Cuculidae, among which it was originally placed, nor in the Primobucconidae; its affinities are at present considered uncertain. Primoscens minutus Harrison and Walker, described in a new Lower Eocene family, Primoscenidae, of Passeriformes, is not a passeriform, but its true affinities are likewise unclear. The presence of the Primobucconidae in the Lower Eocene of England and Wyoming lends support to other evi- dence for a land corridor across the North Atlantic in the early Eocene and further confirms the idea that the small arboreal birds of the Paleogene in the northern hemisphere were non-passerines. The predominant group of small, arboreal birds recorded from the Eocene of North America is the family Primobucconidae, of which two genera and four species are known from the Lower Eocene of Wyoming, and four species in three additional genera from the Middle Eocene, also in Wyoming (Feduccia and Martin, 1976). These birds are apparently most closely related to the modern Bucconidae, which is currently placed in the Piciformes, al- though more probably the Bucconidae, along with the Galbulidae, belong with Coraciiformes (Sibley and Ahlquist, 1972; Burton, 1977; pers. observ.). In a recent study of a Lower Eocene avifauna from England, Harrison and Walker (1977) named a new genus and species of cuckoo (Cuculiformes, Cuculidae), Parvicuculus minor, based on a tarsometatarsus lacking the inner trochlea. Their description and illustrations are sufficient to indicate that this specimen is clearly referable to the Primobucconidae and has no affinity with the Cuculidae. There is precedent for such an error in that the primobucconid Uintornis lucaris Marsh had been referred to the Cuculi- formes (Cracraft and Morony, 1969) until the manifest differences between these two groups were detailed by Feduccia and Martin (1976). We have compared Harrison and Walker’s illustrations and descriptions of Parvicuculus minor with the characters listed in the diagnosis of the Primobucconidae (Feduccia and Martin, 1976:103—104) that are determinable VOLUME 92, NUMBER 3 495 for both Parvicuculus and one or more taxa of primobucconids. Parvicu- culus agrees with the Primobucconidae and differs from the Cuculidae in the following characters: 1) tarsometatarsus short, broad, and flat, not elongate as in the Cuculidae; 2) hypotarsus probably with only a single tendinal canal (Harrison and Walker refer to one hypotarsal groove in Parvicuculus ‘‘which was prob- ably enclosed originally’); 3) papilla for tibialis anticus on the extreme internal margin of the tarso- metatarsus (Harrison and Walker state that this tubercle is ‘‘laterally placed”’ in Parvicuculus but their illustration indicates that this is a lap- sus—if it actually were lateral it would be unlike the condition in the Cuculidae, in which this tubercle is medial but not as much so as in the Primobucconidae); 4) outer trochlea not greatly elevated relative to middle trochlea, not mark- edly expanded or twisted postero-medially to permit full reversal of outer toe. The last feature proves conclusively that Parvicuculus cannot be included in the Cuculidae. ° Harrison and Walker (1977) named a second new genus and species of Lower Eocene Cuculidae, Procuculus minutus, based on an extremely small distal end of a tarsometatarsus. This shows no indication of affinity to the primobucconid Parvicuculus and furthermore bears not the slightest resem- blance to any cuckoo. We are at a loss to understand how this specimen could have been assigned to the Cuculidae and there is nothing by way of explanation in Harrison and Walker’s ‘‘diagnosis,’’ description, or com- ments. The outer trochlea of Procuculus is essentially unmodified and there is no indication that the outer toe was reversed, one of the more diagnostic features of the Cuculidae. The inner trochlea is medially expanded and bears a distinct medially projecting wing which is absent in the Cuculidae. The middle trochlea is decidedly less grooved than in the Cuculidae. Without reference to the actual specimen we are reluctant to say more about Pro- cuculus except that it definitely cannot be referred to either the Cuculidae or the Primobucconidae. The general morphology of the tarsometatarsus of Parvicuculus is similar to that of the Primobucconidae, although the species of that family that are known from comparable three-dimensional tarsometatarsi are all consider- ably larger Middle Eocene forms (and of these the proximal end is known for only one species), so that generic level differences between them and Parvicuculus may be expected. The North American specimens of primo- bucconids that are contemporaneous with Parvicuculus consist of more or less two dimensional, associated skeletons preserved in slabs, so that direct 496 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON comparison of them with Parvicuculus is not possible. Nevertheless, it is of interest that the tarsometatarsus of Parvicuculus minor is almost exactly the same length (11.7 mm) as that of the North American Lower Eocene primobucconid Neanis kistneri (141.5 mm; Feduccia, 1973). Thus there is a possibility that Parvicuculus Harrison and Walker 1977 may prove to be a synonym of Neanis Brodkorb 1965. Regardless of the generic status of Parvicuculus, its significance lies in establishing that the Primobucconidae were present in western Europe at the same time that the family was flourishing in North America. This could be interpreted as supporting the mammalian evidence for faunal interchange between North America and western Europe via a northern corridor across the Atlantic in the early Eocene (McKenna, 1975). The presence of the Primobucconidae in the Old World supports the thesis that the arboreal ‘‘perching’’ birds of the Paleogene were non-passerines (Feduccia, 1977), at least in North America and Europe. For this reason we must remark upon a supposed member of the Passeriformes, also described by Harrison and Walker (1977) from the Lower Eocene of England. This was based on a carpometacarpus which was described as a new genus and species, Primoscens minutus, and which was made the type of a new family of Passeriformes, the Primoscenidae. Harrison and Walker do not say why they assigned this specimen to the Passeriformes, nor do they discuss the relationships of this supposed new family to any of the families or suborders of Passeriformes. One may presume that they were influenced by the fact that the fossil has a fairly well developed intermetacarpal tuberosity, but among the ‘‘subpasserine’’ land birds this feature also occurs in the Coli- iformes, Piciformes, and some of the Coraciiformes. At least one character used in the diagnosis of Primoscens (‘‘distal facet for digit 3 extends only a little beyond that of digit 2’), indicates that the genus is misplaced as to order, for even in the most primitive passerines the third metacarpal char- acteristically extends distally well beyond the second metacarpal. We note as much similarity between Primoscens and some of the primitive Coraci- iformes, such as Brachypteracias (Brachypteraciidae), as with the Passer- iformes. We do not consider that Primoscens can be regarded as a passer- ine, and as yet there is no undoubted passeriform fossil of Eocene age known (cf. Brodkorb, 1978). Acknowledgments For reading and commenting on the manuscript we are grateful to Pierce Brodkorb, John Farrand, Jr., and David W. Steadman. Literature Cited Brodkorb, Pierce. 1965. New taxa of fossil birds.—Quarterly Journal of the Florida Academy of Science 28:197-198. VOLUME 92, NUMBER 3 497 . 1978. Catalogue of fossil birds. Part 5 (Passeriformes).— Bulletin of the Florida State Museum, Biological Sciences 23(3):139—228. Burton, Philip J. K. 1977. Feeding behavior in the Paradise Jacamar and the Swallow-wing.— Living Bird 15(‘‘1976’’):223-238. Cracraft, J., and J. J. Morony. 1969. A new Pliocene woodpecker, with comments on the fossil Picidae —American Museum Novitates 2400: 1-8. Feduccia, Alan. 1973. A new Eocene zygodactyl bird.—Journal of Paleontology 47(3):501- 503. . 1977. A model for the evolution of perching birds.—Systematic Zoology 26(1):19-31. , and L. D. Martin. 1976. The Eocene zygodactyl birds of North America (Aves: Piciformes).—Smithsonian Contributions to Paleobiology 27:101-110. Harrison, C. J. O., and C. A. Walker. 1977. Birds of the British Lower Eocene.—Tertiary Research Special Paper 3:1—52, 11 plates. McKenna, Malcolm. 1975. Fossil mammals and early Eocene North Atlantic land continui- ty.—Annals of the Missouri Botanical Garden 62(2): 335-353. Sibley, C. G., and J. E. Ahlquist. 1972. A comparative study of the egg white proteins of non- passerine birds.—Peabody Museum of Natural History Yale University Bulletin 39:1- 276: (SLO) Department of Vertebrate Zoology, National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560, and (AF) De- partment of Zoology, University of North Carolina, Chapel Hill, North Car- olina 27514. PROC. BIOL. SOC. WASH. 92(3), 1979, pp. 498-504 A NEW SPECIES OF ELEUTHERODACTYLUS FROM NORTHERN ECUADOR (AMPHIBIA: LEPTODACTYLIDAE) John D. Lynch Abstract.—Eleutherodactylus loustes, new species, is described from in- termediate elevations on the Pacific versant of the Andes in northern Ec- uador. The new species exhibits a combination of character states inter- mediate between those of frogs of the fitzingeri and unistrigatus groups and differs from all congeners in having toe webbing, areolate skin on the venter, and the first finger longer than the second. Its closet relatives are E. cren- unguis Lynch and E. latidiscus (Boulenger) from the Pacific versant and E. cremnobates Lynch and Duellman and E. rubicundus (Jiménez de la Es- pada) from the Amazonian versant. Lynch (1976a) defined the species groups of Eleutherodactylus in South America primarily on the basis of relative lengths of the inner two fingers and the texture of the skin on the venter. The two largest species groups he recognized in South America were the fitzingeri group (ca. 40 species) and the uwnistrigatus group (ca. 100 species). Lynch (1976b) and Lynch and Duellman (1979) named two species which strained those group definitions. Eleutherodactylus crenunguis was placed in the fitzingeri group by Lynch (1976b) who noted its resemblance to E. cruentus (Peters) and E. latidiscus (Boulenger). Lynch and Duellman (1979) removed E. crenunguis to the unistrigatus group, named E. cremnobates, and considered these two species closely related to one another as well as to E. rubicundus and E. latidiscus; the four species were placed in the rubicundus assembly within the uwnistrigatus group. During field work in northern Ecuador in May 1977, an additional species of this complex was found. It differs trenchantly from the four species known to date in having appreciable toe webbing. The following abbreviations are employed: SVL (snout-vent length), IOD (interorbital distance), E-N (eye to nostril distance), and KU (University of Kansas Museum of Natural History). Eleutherodactylus loustes, new species (Figs. 1, 2) Holotype.—KU 179234, an adult female, one of a series collected at Mal- donado, Provincia Carchi, Ecuador, 1410 m, on 29 May 1977 by Thomas Berger, David Cannatella, and John Lynch. VOLUME 92, NUMBER 3 499 Fig. 1. Eleutherodactylus loustes (KU 179233), adult male, 31.2 mm SVL; from Koda- chrome by author. Paratypes.—KU 179231-33, 179235—50, collected at the type locality on 29 and 30 May 1977. Diagnosis.—1) skin of dorsal surfaces smooth except for anastomosing ridgelets on lower back and prominent fold between eyes, that of venter areolate; no dorsolateral folds; 2) tympanum concealed beneath skin, an- nulus small; 3) snout round in dorsal view, angularly rounded in lateral profile; 4) upper eyelid broader than IOD, bearing low warts; low cranial crests palpable; 5) vomerine odontophores elevated, oval; 6) males with vocal slits and subgular vocal sac; no nuptial pad on thumb; 7) first finger slightly longer than second; fingers II-IV bearing enlarged pads; all pads bearing broad discs; 8) fingers bearing distinct lateral fringes; 9) no ulnar tubercles except small antebrachial; 10) no tubercles on heel or tarsus; flap- like inner tarsal fold on distal 7s—’%2 of tarsus; 11) two metatarsal tubercles, inner oval, 8 times size of subconical outer; 3—6 indistinct supernumerary plantar tubercles; 12) toes bearing prominent flap-like lateral fringes, webbed—modal formula I 2~-—2 II 14%4—3- HI 2*—3% IV 3%—2* V; fringe along outer edges of toes I and V; toe pads smaller than those of outer fingers; 13) brown above with darker brown markings on occiput and sacrum; labial and limb bars, canthal-supratympanic stripes dark brown; anterior and posterior surfaces of thighs brown; throat gray; venter cream with diffuse brown reticulation; 14) adults large, males 31.2— 37.1 @ = 34.8, N = 6), one adult female 46.7 mm SVL. Eleutherodactylus loustes is distinguished from its nearest relatives (E. cremnobates, E. crenunguis, E. latidiscus and E. rubicundus) by virtue of its toe webbing and inner tarsal fold. Description.—Head as wide as (or wider than) body; head wider than long; head width 38.0-40.9 @ = 39.5, N = 10) percent SVL; snout suba- 500 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 2. Eleutherodactylus loustes. A, B, dorsal and lateral views of head, KU 179242; C, ventral view of foot, KU 179243. Line equals 5 mm. cuminate to round in dorsal view, angularly rounded in lateral profile; nos- trils weakly protuberant, directed dorsolaterally; snout short, E-N of males 67.8-81.6 (* = 74.8, N = 6) percent eye length, of females 85.2-91.5 @ = 88.7, N = 4) percent; canthus rostralis evident but rounded, weakly con- cave (Fig. 2); loreal region concave, gently sloping to lips; lips not flared; interorbital space narrow, bearing an interocular ridge with a median tu- bercle; cranial crests palpable; upper eyelid width 100.0-150.0 @ = 121.5, N = 10) percent IOD; low, non-elevated tubercles on upper eyelids; thick supratympanic fold from posterior corner of eye to above insertion of upper arm; postrictal tubercles large, elevated; tympanic annulus concealed be- neath thick skin of side of head (Fig. 2); choanae large, oval, longer than wide, not concealed by palatal shelf of maxillary arch when roof of mouth is viewed from directly below; vomerine odontophores elevated, oval, me- VOLUME 92, NUMBER 3 501 dian and posterior to choanae, separated by a distance equal to between % and a full width of an odontophore, each slightly larger than a choana, bearing a transverse row of 4—5 teeth in males, 7-8 teeth in large females; tongue large, as long as wide, posterior edge deeply notched, posterior '/s not adherent to floor of mouth; males having long vocal slits posterolateral to tongue; vocal sac external, subgular, single. Skin of dorsal surfaces smooth except for small anastomosing ridges; vent extended in a sheath to level of posterior edge of thigh but not going down on to back of thighs; skin of venter areolate; throat smooth except for small, white warts; forearm bearing small antebrachial tubercle, but no other ulnar tubercles; palmar tubercles low; palmar tubercle bifid, twice as large as oval thenar tubercle; supernumerary palmar tubercles large, diffuse; subarticular tubercles low, round; fingers bearing keel-like lateral fringes and large pads (2-3 times width of digit below pad) bearing broader than long discs on fingers II-IV; pad of thumb small but bearing broad disc; all pads and discs apically rounded (not emarginate); male lacking nuptial pad on thumb; first finger longer than second. Heel lacking tubercles or bearing one indistinct tubercle; outer edge of tarsus lacking tubercles; inner edge of tarsus bearing strong (flap-like) inner tarsal fold along distal 2/s to % of tarsus; inner metatarsal tubercle flat, oval, outer metatarsal tubercle subconical, ’% size of inner; plantar supernumerary tubercles indistinct (if present at all); subarticular tubercles not elevated; prominent fringe along inner edge of toe I and lesser fringe along outer edge of toe V; toe pads slightly smaller than those of outer fingers; toes basally webbed (Fig. 2), free portions of toes bearing very prominent lateral fringes; webbing formulae (following Savage and Heyer, 1967) for males I (2—27-)— (2+-2) HT 1%-1%)— (3-27) HI (244-2)— (374-344) IV (342-34%4)—(2"s—2) V, for females I 2-—(2*-2) I] (%-— 1%)—(%-3-) HI (2t-2-) —G%-3%) IV 344—(2t-2-) V; shank of males 56.8-63.1 @ = 59.2, N = 6) percent SVL, of females 58.8-61.3 @ = 60.4, N = 4) percent. Ground color brown; body bearing dark brown occipital W-shaped mark, interorbital bar, broad sacral chevron (almost a band); occipital W bearing indistinct ridges outlined with cream; interorbital bar bearing cream line along anterior border; sacral chevron outlined in cream; dark bars on thighs brown separated by pale brown interspaces which are subdivided by dusky brown lines; posterior surfaces of thighs dusky brown; dark brown bars on shanks as wide as (or slightly narrower than) pale brown interspaces, shank bars more or less perpendicular to limb axis; canthal-supratympanic stripe and labial bars dark brown; indefinite slanted bars on flanks; anterior sur- faces of thighs brown with small cream spots; throat gray with white flecks; venter cream with diffuse brown reticulation; throat and venter pigmenta- tion separated by diffuse brown V-shaped mark; undersides of limbs gray flecked with white and bearing diffuse brown reticulation. 502 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON In life, E. loustes is colored as follows: ‘‘Large female: reddish brown with black spots. Some green wash on spaces between leg bars and on lower flank. Concealed thigh black with greenish speckles. Iris deep brown. Ven- ter cream with brown spots. Throat brown with cream flecks. Toe webs pale yellow-green. Male: green with gold or copper wash, especially ante- riorly. Iris more coppery [otherwise like female]’’ (J. D. Lynch fieldnotes 29 May 1977). Measurements of holotype in mm.—SVL 46.7; shank 28.5; head width 19.1; head length 16.1; upper eyelid width 4.5; IOD 3.3; eye length 6.0; E- INGS-3 Natural history.—Specimens were collected on the nights of 29 and 30 May 1977 on bare rock faces, moss encrusted rocks on the floor of the stream, and on leaves in the spray zone of a waterfall. The small stream (a tributary of the Quebrada Naranjo) was heavily shaded by forest. When approached with a headlamp the frogs crouched against the wet substrate. Collecting one hundred meters on either side of the small waterfall failed to yield additional specimens nor were specimens encountered in seemingly similar microhabitats along the nearby Quebrada Huagambi (both debauch into the Rio San Juan within 200 m). No evidence of reproductive activity was noted although males have swollen testes. Two females (JDL 8641-42) are slightly smaller (44.9 and 46.0 mm SVL) than the holotype and have feebly convoluted oviducts. Etymology.—Greek, meaning one fond of bathing. The name is in loose allusion to the frogs being found only within the spray zone of a waterfall in a deep, cool canyon. Sympatric congeners.—In addition to E. loustes, E. achatinus (Boulen- ger), E. chalceus (Peters), E. calcarulatus Lynch, E. necerus Lynch, and E. w-nigrum (Boettger) were secured in five nights of fieldwork (27-31 May 1977). One other species found at Maldonado is being described separately (Lynch and Myers, MS). None of these were found in the same microhabitat as E. loustes. Discussion.—Lynch and Duellman (1979) suggested that E. crenunguis (Pacific) and E. rubicundus (Amazonian) were a species pair in the rubi- cundus assembly and that E. cremnobates (Amazonian) and E. latidiscus (Pacific) were a second pair. Eleutherodactylus loustes is much more similar to E. crenunguis and E. rubicundus than to E. cremnobates or E. latidiscus. All five species share the following character states: interorbital space narrow, cranial crests present (usually adult females only), fingers and toes long and slender and bearing large pads (and broad discs); vomerine odon- tophores prominent; canthus rostralis obsolete (not sharp); loreal region sloping gently to lips. The tympanic annulus is small in all species but is concealed beneath the skin only in E. loustes. That species also differs from VOLUME 92, NUMBER 3 503 Table 1. Character states in the five species of the rubicundus assembly of the uwnistrigatus group of Eleutherodactylus (X = yes, O = no). cremnobates crenunguis latidiscus loustes rubicundus Male with vocal slits O > O x First finger > second O x O x x Venter finely areolate x O x x x Digital pads emarginate O x x O x Heel and tarsal tubercles x x O O x Lateral fringes on fingers x O x x O the others in having toe webbing, prominent toe fringes, and a flap-like inner tarsal fold. All five species are nocturnal. Eleutherodactylus cremnobates, E. loustes, and E. rubicundus are found in very wet areas (often spray zones or drip zones) along small rocky streams (0.5—1.5 m wide) in dense cloud forests. Eleutherodactylus crenunguis is sometimes encountered in such situations but is more frequently found in forest-edge microhabitats. Eleutherodactylus latidiscus is a lowland forest species and specimens have accumulated slow- ly. The apparent rarity may reflect the frogs’ preference for a higher stratum in the forest than that normally searched. Richard Zweifel obtained a calling male 4 meters above ground on ‘‘a 6” diameter partly hollow and branchless snag, laden with bromeliads.”’ Those characteristics readily reduced to discrete character states (Table 1) do not obviously correlate with the ecological notes now available for the five species (nor do they readily correlate with my views on the relationships among the five species). However, neither do they sort the taxa geograph- ically. Collections of eleutherodactyline frogs in Ecuador are extensive and al- though much data remain to be published (including approximately 25 un- described species), no other eleutherodactyline species are known to me that might be closely related to this assembly of five species. A critical point is the absence of rubicundus assembly frogs at elevations above 1700 me- ters. The reported distribution of E. latidiscus covers eastern Panama, west- ern Colombia, and northwestern Ecuador (Cochran and Goin, 1970) but the Panamanian material as well as much of the Colombian material is misiden- tified. I have examined E. latidiscus as far north as the drainage of the Rio San Juan in Colombia and as far south as the Rio Palenque Biological Station in southern Provincia Pichincha in Ecuador. Cochran and Goin’s (1970) E. latidiscus tamsitti, if distinct, establishes the occurrence of the species in the headwater regions of the Rio Magdalena. The presence of two sets of species found in low cloud forests on opposite 504 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON sides of the Ecuadorian Andes where each set has trans-Andean and cis- Andean representatives poses a clue to the role of the Andes in the im- pressive speciation of Eleutherodactylus in northwestern South America. The distributional and phylogenetic patterns suggest that the Andean orog- eny isolated the crenunguis-loustes and the latidiscus stocks from those of rubicundus and cremnobates respectively. Acknowledgments.—Specimens were loaned by William E. Duellman and Richard G. Zweifel. Duellman, Alice Grandison, Ron Heyer, and Zweifel provided working space and other amenities at their respective institutions. Travel to museums was supported by the University of Nebraska Research Council. Funds for fieldwork were provided by a grant from the American Philosophical Society and a Summer Research Fellowship (University of Nebraska Research Council). Thomas Berger and David Cannatella aided me in the field. Literature Cited Cochran, D. M., and C. J. Goin. 1970. Frogs of Colombia.—U.S. Nat. Mus. Bull. 288, 655 pp. Lynch, J. D. 1976a. The species groups of the South American frogs of the genus Eleuther- odactylus (Leptodactylidae)—Occ. Pap. Mus. Nat. Hist. Univ. Kansas 61:1—24. . 1976b. New species of frogs (Leptodactylidae: Eleutherodactylus) from the Pacific versant of Ecuador.—Occ. Pap. Mus. Nat. Hist. Univ. Kansas 55:1-33. Lynch, J. D., and W. E. Duellman. 1979. The Eleutherodactylus of the Amazonian slopes of the Ecuadorian Andes (Anura: Leptodactylidae).—Misc. Publ. Univ. Kansas Mus. Nat. Hist. in press. Savage, J. M., and W. R. Heyer. 1967. Variation and distribution in the tree-frog genus Phyllomedusa in Costa Rica, Central America.—Beit. Neotrop. Fauna 5:111—-131. School of Life Sciences, The University of Nebraska, Lincoln, Nebraska 68588. PROC. BIOL. SOC. WASH. 92(3), 1979, pp. 505-509 PEOSIDRILUS BIPROSTATUS N. G., N. SP., A MARINE TUBIFICID (OLIGOCHAETA) FROM THE EASTERN UNITED STATES H. R. Baker and Christer Erséus Abstract.—A new monotypic genus Peosidrilus is established for the marine tubificid P. biprostatus n. sp. from New Jersey and North Carolina. The species is characterized by its possession of cylindrical atria ending in large, protrusible penes, two pairs of pedunculate prostate glands, paired spermathecae, and bundles of penial setae. Peosidrilus appears closely re- lated to Phallodrilus Pierantoni, 1902. A new tubificid oligochaete was found at several stations on the conti- nental shelf off New Jersey, in the course of a Bureau of Land Management baseline study made by staff at the Virginia Institute of Marine Science (principal investigator, Dr. D. Boesch). An additional specimen was found in coastal water of North Carolina, during a stay at the Duke University Marine Laboratory, Beaufort, N.C. The tubificid represents a new mono- typic genus within the sub-family Phallodrilinae Brinkhurst, 1971. Material and Methods Specimens of P. biprostatus n. g., n. sp. from New Jersey were placed at our disposal by Dr. R. J. Diaz, Virginia Institute of Marine Science. Most of the worms were stained in Fast Green FCF, a few others in paracarmine. The microscopical studies were then made on either dissected or whole- mounted individuals. The specimen from North Carolina was fixed in Bouin’s fluid, sectioned and stained in Heidenhain’s haematoxylin. The type-series of the new species is deposited in the United States Na- tional Museum of Natural History (USNM), Smithsonian Institution, Wash- ington, D.C. Peosidrilus, new genus Type-species.—Peosidrilus biprostatus n. sp., by original designation. Etymology.—Worm (‘drilus’ Greek) with penis (‘peos’ Greek). Gender masculine. Definition.—Marine tubificids. Hair setae absent. Penial setae present ventrally in XI. Spermathecal setae absent. Male and spermathecal pores paired in XI and X respectively. Coelomocytes, if present, small and sparse, not of the ‘Rhyacodriline-type.’ 506 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON A B C Fig. 1. Peosidrilus biprostatus: A, Ectal tip of anterior somatic seta; B, Ectal tip of posterior somatic seta; C, Ectal tip of penial seta. Vasa deferentia ciliated, entering apical, ental ends of atria. Atria cylin- drical and ciliated, each bearing 2 pedunculate prostate glands, one at ental, one at ectal end. Atria entering, but separated by constriction from, large penes enclosed in deep penial sacs. Spermathecae paired, filled with random masses of sperm. Spermatophores not developed. Peosidrilus biprostatus, new species Figs. 1—2 Holotype.—USNM 57074, a whole-mounted specimen. Type-locality.—Continental shelf off New Jersey, U.S.A., 39°29.9’N, 73°10.1'W, 41 m, coarse sand (type date: Aug. 1976). Paratypes.—USNM 57075-57078. One whole-mounted spm from the type-locality. Three whole-mounted and 4 dissected spms from 39°15.2’'N, 74°09.2'W, 24 m, coarse sand (Nov. 1975). One whole-mounted spm from 39°15.2'N, 74°09.1’W, 25 m, coarse sand (June 1976). Sections of genital region of one spm from Beaufort area, North Carolina: at Shackleford Point, E of buoy N"2”, 34°41'14"N, 76°38'30’”W, 5.5 m, coarse sand with some shell gravel (Amphioxus-sand) (Oct. 25, 1977; C. E. coll.). Other material.—In R. O. Brinkhurst collection: 5 spms from various stations off New Jersey, depths ranging from 17 to 73 m, mostly coarse sands. In C. E. collection: 15 spms from same area, depths from 16 to 63 m, coarse or medium sands. Description.—Length (5 spms) 6.9-9.5 mm, 54—62 segments. Prostomium rounded, about as long as broad. Clitellum well developed on 2 X—XII of fully mature specimens. Somatic setae (Fig. 1A—B) bifid, with upper teeth much thinner and shorter than lower (upper teeth become somewhat broader pos- teriorly; Fig. 1B). Bifids 3—4 per bundle throughout body, up to 47 um long in V—VI, in other parts of body slightly shorter. Ventral setae of XI modified into penial bundles, each of which contains 7—13 straight, but ectally strong- VOLUME 92, NUMBER 3 507 508 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON ly hooked, penial setae (Fig. 1C). Penials 30-37 um long (may vary within bundle), about 2.5 wm thick. Male pores paired, located in line with ventral setae in posterior part of XI. Spermathecal pores paired, located laterally in most anterior part of X, very near intersegmental furrow IX/X. Pharyngeal glands in IV—VI. Male genital system (all structures paired) (Fig. 2): vas deferens (vd) ciliated, slender (6-8 um thick), longer than atrium; vas deferens entering apical, ental end of cylindrical atrium; atrium (a), 90-140 um long, 27-37 wm wide, with outer muscular lining (very slen- der in ental part, up to 5 wm thick in ectal part of atrium), and inner ciliated epithelium, which is densely granulated in ental and in dorsal-middle part of atrium; ventral-middle and ectal parts of atrial epithelium generally non- granulated, but consisting of very large, apparently vacuolized cells, one large prostate gland (pr /) attached by stalk to ental, sub-apical part of atrium, near entrance of vas deferens; a posterior large prostate (pr 2) con- nected with dorsal middle-to-ectal part of atrium by means of a very long stalk; atrium separated from large, pear-shaped, 80-120 um long, penis (p) by constriction; penis basally bulbous and about 30-40 um wide, ectally tapering with a slightly bulbous tip, 12-16 wm wide, internal epithelium of penis ciliated; epithelium with large, vacuolized cells in basal part of penis; penis enclosed in deep penial sac (psa), into which also protrude ectal tips of penial setae (pse); cuticular penial sheath absent. Spermathecae (Fig. 2, s) paired and cylindrical, 110-170 wm long and generally not wider than 25— 40 um, divided into short, but distinct, thick-walled ducts, and thin-walled ampullae. Spermathecal ampullae filled with random masses of sperm. Habitat.—Subtidal, coarse to medium sands. Distribution.—New Jersey and North Carolina, U.S.A. (NW Atlantic). Discussion Peosidrilus is clearly a phallodriline genus, and it appears most closely related to Phallodrilus Pierantoni, 1902, which also is characterized by hav- ing more or less cylindrical atria with 2 pairs of prostate glands. The penial setae of P. biprostatus are almost identical to those of, for instance, Phal- lodrilus coeloprostatus Cook, 1969, and Phallodrilus prostatus (Knollner, 1935). However, the present definition of Phallodrilus (Erséus, in press) does not include species with true penes, organs that are very conspicuous in the new species. It is not considered appropriate to extend Phallodrilus to include biprostatus, but to establish the new genus Peosidrilus for it. Aktedrilus Kn6llner, 1935, is another phallodriline genus with 2 pairs of prostates and true penes. However, all its forms are characterized by un- paired spermathecae and the complete absence of penial setae (Finogenova, 1976; Erséus, in preparation). The development of penes in both Aktedrilus and Peosidrilus might well be just a matter of convergence. VOLUME 92, NUMBER 3 509 The histological differentiation in the different parts of the atrium of P. biprostatus could be taken as an indication of a close phylogenetic relation- ship to Spiridion Knollner, 1935, which has been reviewed separately (Erséus, 1979). The atrium of Spiridion consists of a granulated ental part and a non-granulated ectal part, the latter of which has a winding lumen, and which can be everted during copulation at least in S. insigne Knollner, 1935. However, Spiridion possesses neither true penes nor posterior pros- tates. Acknowledgments We are very much indebted to Dr. R. J. Diaz, VIMS, Gloucester Pt., Va., for sending us oligochaete material. H. R. B. would like to thank Dr. R. O. Brinkhurst for his valuable criticism and encouragement. C. E. would like to thank Dr. J. D. Costlow for working facilities at the Duke Marine Lab- oratory, Beaufort, N.C.; and to the Duke University, the Commonwealth of Virginia, and the Swedish Natural Science Research Council, for financial support. Literature Cited Erséus, C. 1979. Re-examination of the marine genus Spiridion Knollner (Oligochaeta, Tubificidae).—Sarsia 64: 183-187. In press. Taxonomic revision of the marine genus Phallodrilus Pierantoni (Oli- gochaeta, Tubificidae), with descriptions of thirteen new species.—Zool. Scripta. Finogenova, N. P. 1976. New species of Tubificidae (Oligochaeta) from the Caspian Sea. (In Russian.)—Zool. Zh. 55:1563—1566. (HRB) Ocean Ecology Laboratory, Institute of Ocean Sciences, Patricia Bay, Sidney, British Columbia, Canada V8L 4B2; (CE) Department of Zo- ology, University of Gothenburg, Fack, S-400 33 Goteborg, Sweden PROC. BIOL. SOC. WASH. 92(3), 1979, pp. 510-526 NEW GASTROPODS FROM THE ABROLHOS ARCHIPELAGO AND REEF COMPLEX, BRAZIL Edward J. Petuch Abstract.—Ten new gastropods, Acmaea (Collisella) abrolhosensis, Cy- phoma macumba, Dermomurex (Trialatella) oxum, Muricopsis oxossi, Mu- rexiella iemanja, Latirus (Polygona) ogum, Oliva (Plicoliva) zelindae, Vex- illum (Costellaria) kaicherae, Vexillum (Costellaria) lixa, and Conus iansa, are described from the Abrolhos Archipelago and reef complex, Bahia State, Brazil. A new subgenus, Plicoliva, of the genus Oliva, family Olividae, is also described. The unusual nature of the Abrolhos mollusk fauna and the unique ecological conditions characteristic of the area are outlined. The Abrolhos Archipelago region (ranging from approximately 17°26’S to 18°11’S and 38°34’W to 39°17'W) off the south coast of Bahia State, Brazil, comprises 5 islands, 2 major coral reef complexes, and innumerable smaller isolated reefs. The archipelago (approx. 17°57’S, 38°42’W) includes the is- land of Santa Barbara, about 2 km in length, and the smaller islands Siriba, Redonda, Sueste, and Guarita, and represents the southernmost extensive coral reef growth in the western Atlantic (Laborel, 1969: 125, fig. 45). The Abrolhos area is malacologically poorly known; only small, incom- plete collections have been taken since Charles Darwin briefly visited the archipelago in the 1830’s. Hartt (1870: 174-214) published an account of the geology and natural history of the Abrolhos reefs and islands but only men- tioned a few species of mollusks. In this century, the only major expedition to collect around the islands was that of the CALYPSO in 1961-1962. This involved a few days SCUBA diving on some of the adjacent reef complexes and was by no means comprehensive. During July, 1977, I had the oppor- tunity to visit the archipelago and surrounding reefs. Material was acquired by SCUBA diving, snorkeling, and reef and shore collecting at low tide. Studies of this material showed that several gastropods were new to science; the new taxa are described herein. Physical Characteristics of the Abrolhos Area The Abrolhos Archipelago are high islands and have coastlines composed of steep cliffs with low bordering rock platforms. Geologically, they are a mixture of sedimentary and igneous rocks (Hartt, 1870: 174-176) and their history has still not been completely worked out. Between the mainland and VOLUME 92, NUMBER 3 S11 the islands there are extensive coral reef complexes—the largest of these being the rectangular-shaped Parcel das Paredes, a reef of over 30 km length on a side. North and south of the Parcel das Paredes are series of large reefs that are exposed at low tide. Another extensive reef system, the Parcel dos Abrolhos, borders the edge of the continental shelf just east of the archi- pelago. Scattered throughout the area are large numbers of small (10-50 m di- ameter) vase or inverted wide-brimmed hat-shaped reefs called *‘chapei- roes’’ by the native Bahianos. These reefs, composed primarily of 3 species of Mussismillia corals, Millepora hydrocorals, and coralline algae, rise abruptly from the flat sea floor of the Abrolhos Platform. These giant ‘‘hats”’ spread horizontally just below the sea surface and eventually fuse with other ‘‘chapeiroes’’ to form the larger reef complexes (Laborel, 1969: 136-137, figs. 51-52). The sea floor between the mainland and the ‘“‘chapeiroes,”’ reef complexes, and high islands is composed primarily of soft calcareous muds and oozes. Mean depth throughout the Platform area is only about 25 m. Aspects of the Abrolhos Gastropod Fauna New species described in this paper include: Family Patellidae Acmaea (Collisella) abrolhosensis n. sp. Family Ovulidae Cyphoma macumba n. sp. Family Muricidae Dermomurex (Trialatella) oxum n. sp. Muricopsis oxossi Nn. sp. Murexiella iemanja n. sp. Family Fasciolariidae Latirus (Polygona) ogum N. sp. Family Olividae Oliva (Plicoliva) zelindae n. subgen. and n. sp. Family Costellariidae Vexillum (Costellaria) kaicherae n. sp. Vexillum (Costellaria) lixa n. sp. Family Conidae Conus iansa Nn. sp. Besides these new species, several other Abrolhos gastropods are noted in order to emphasize the unusual aspects of the fauna. These are arranged by ecological preferences. 1. Rocky Shorelines of the Islands.—In the supratidal zone, only 3 species of Littorina were collected. Littorina ziczac (Gmelin, 1791) and L. 512 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON flava King and Broderip, 1832, were rare, with only 2 specimens of each taken on Santa Barbara. On the other hand, L. angulifera (Lamarck, 1822) (the mangrove periwinkle) was abundant even though there were no man- groves on the Abrolhos. These were much larger, heavier, and stockier than the typical form. They were also more colorful, often with dark spottings of purple and red. At the mean low water line, only Leucozonia brasiliana (d’Orbigny, 1841), Leucozonia lineata Usticke, 1969, Acmaea (Collisella) abrolhosensis n. sp., and Siphonaria hispida E. A. Smith, 1890, were common. 2. Fringing Reefs and ‘‘Chapeiroes.’’—At depths of 2 m and less on the reef platforms, commonly encountered species included Vasum cassiforme (Kiener, 1841), Pleuroploca aurantiaca (Lamarck, 1816), Latirus (Poly- gona) ogum n. sp., Cerithium atratum (Born, 1778), Pisania pusio (Lin- naeus, 1758), Thais rustica (Lamarck, 1822), and Astraea latispina (Philip- pi, 1844). Fresh dead specimens of Cypraea zebra dissimilis Schilder, 1924, and Conus brasiliensis Clench, 1942, were frequently found on these reefs. 3. Shallow Water Sand-Bottom Areas.—Scattered among the larger reefs were pockets of coarse sand and coral rubble. These often accumulated in ‘‘bays’’ at the edges of the large eroded reef complexes. Species frequently encountered here were Turbinella laevigata Anton, 1839, Cassis tuberosa (Linnaeus, 1758), Oliva circinata Marrat in Sowerby, 1871, Oliva (Plicoliva) zelindae n. subgen. and n. sp., Vexillum (Costellaria) kaicherae n. sp., Closia largillieri (Kiener, 1841), Olivella verreauxi (Duclos, 1857), and Co- nus jaspideus Gmelin, 1791. 4. Carbonate Mud Bottom.—At depths of 20-25 m, the calcareous ooze substrate was covered with a thin layer of green algae. Crawling through this algal mat were Conus iansa n. sp., Nassarius capillaris (Watson, 1882), Mitrella albovittata Lopes, Coelho, and Cardoso, 1965, Turritella exoleta (Linnaeus, 1758), Bullata bullata (Born, 1778), and Typhis cf. sowerbyi (Broderip, 1833). | In general, gastropod species diversity was much lower than could be expected from an area of such diverse habitats. Though the species endemic to the islands are interesting, the absence of characteristic tropical western Atlantic rocky intertidal genera such as Nerita, Tectarius, Echininus, and Nodilittorina is especially noteworthy. Systematic Part Abbreviations used in this paper: USNM—United States National Mu- seum Collection (now National Museum of Natural History); UMML—In- vertebrate Museum Collection of the Rosenstiel School of Marine and At- mospheric Science, University of Miami; DMNH—Delaware Museum of Natural History. VOLUME 92, NUMBER 3 513 Family Patellidae Genus Acmaea Rathke in Eschscholtz, 1833 Subgenus Collisella Dall, 1871 Acmaea (Collisella) abrolhosensis, new species Fig. 1, A and B Material Examined.—HOLOTY PE—Length 22 mm, width 19 mm, Santa Barbara Is., Abrolhos Archipelago (17°58'S, 38°42'W), 27 July 1977, USNM 780644. PARATYPES—Lengths 21 mm, 18 mm, 17 mm, 17 mm, USNM 780645; lengths 21 mm, 21 mm, 20 mm, 19 mm, DMNH 121797; lengths 18 mm, 15 mm, 14 mm, Museu Nacional do Brasil, Rio de Janeiro, 3733; lengths 26 mm, 22 mm, 15 mm, 14 mm, 15 mm, 14 mm, 13 mm, UMML 8163-8169; all specimens from same locality, depth, and date as holotype. Shell Description.—Shell flattened, smooth, oval in outline, fairly thick for its size; apex pointed with sloping sides; external color black with wide white bands radiating from apex; interior cream-colored with dark external radial color pattern showing through; callus dark brown with bluish-white center; edge of shell scalloped with undulating margin conforming to radial color pattern. Coloration of Animal.—Pale greenish in life, fading to creamy white when preserved in ethyl alcohol. Type-locality.—On rock, low water line, south side of Santa Barbara Is., Abrolhos Archipelago, Bahia State, Brazil (19°57'S, 38°42’W). Distribution.—The new species is endemic to the rocky coasts of the islands of Santa Barbara, Redonda, Siriba, Sueste, and Guarita, of the Abrolhos Archipelago. Ecology.—Acmaea abrolhosensis is acommon species on all of the Abrol- hos Islands. It is found primarily on algae-covered rocks at and slightly below the mean low tide line. The new species is most often associated with the gastropods Siphonaria hispida E. A. Smith, 1890, and Littorina angulif- era (Lamarck, 1822). Etymology.—Named for the Abrolhos Islands. Remarks.—The shell of A. abrolhosensis is highly variable in color; some specimens are almost pure white while others are almost completely black. These darker morphs resemble the Fernando de Noronha Is. endemic, A. noronhensis E. A. Smith, 1890 (E. A. Smith, 1890:475—520, Righi, 1966: 270, figs. 1-2, Rios, 1975:pl. 4, fig. 40). That species, however, lacks the white radiate markings that are characteristic of A. abrolhosensis. The clos- est relative of the new species is A. subrugosa d’Orbigny, 1846; this species ranges from Alagoas, Brazil, south to Uruguay. A. subrugosa differs from A. abrolhosensis by having a consistently finer external white radiating pattern, by having a larger central muscle scar, and by lacking the bluish- white clouding at the center of the callus. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON. Fig. 1. A, Acmaea (Collisella) abrolhosensis, dorsal aspect of holotype; B, Acmaea (Col- lisella) abrolhosensis, ventral aspect of holotype; C, Cyphoma macumba, ventral aspect of holotype; D, Cyphoma macumba, dorsal aspect of holotype; E, Dermomurex (Trialatella) oxum, dorsal aspect of holotype; F, Dermomurex (Trialatella) oxum, ventral aspect of holo- type; G, Muricopsis oxossi, dorsal aspect of holotype; H, Muricopsis oxossi, ventral aspect of holotype; I, Murexiella iemanja, dorsal aspect of holotype; J, Murexiella iemanja, ventral aspect of holotype. VOLUME 92, NUMBER 3 515 Acmaea abrolhosensis is the newest member of an interesting species complex. This includes the mainland A. subrugosa and 3 offshore species, each endemic to a different island group; A. abrolhosensis to the Abrolhos Archipelago, A. noronhensis to Fernando de Noronha Is., and A. marcusi Righi, 1966, to Trindade Is. (Righi, 1966:269-270, figs. 1-6). The offshore island forms, though morphologically distinct, are similar to A. subrugosa and probably represent recent offshoots from that species. The radulae of the 4 species differ greatly from each other. Righi (1966:277, figs. 12-14) illustrates and discusses the differences in radulae between A. subrugosa, A. noronhensis, and A. marcusi. When the radula of A. abrolhosensis (Fig. 2A) is compared with those of the 3 other species, it can be seen to be intermediate between A. subrugosa and A. noronhensis. However, the extremely well-developed cusp on the second lateral tooth is characteristic of the new species and sets it aside as distinct. Family Ovulidae Genus Cyphoma Roding, 1798 Cyphoma macumba, new species Fig. 1, C and D Material Examined.—HOLOTYPE—length 22 mm, width 13 mm, 2 m depth off Parcel das Paredes Reef, Abrolhos Reef Complex (17°47'S, 39°1’W), 24 July 1977, USNM 780646. PARATYPE: length 30 mm, Lixa Reef, Parcel das Paredes Reef area, Abrolhos Reef Complex (17°43’S, 38°59'W) 23 July 1977, USNM 780647. Shell Description.—Elongate, shiny, with fine longitudinal striations; dor- sal ridge prominent, sharp; outer lip thickened with protruding hornlike projection corresponding to dorsal ridge; color pale cream-orange to white; interior of aperture pale orange. . Animal.—Mantle transparent, with 4 groups of roughly rectangular- shaped markings arranged in longitudinal rows (Fig. 2B); rectangular mark- ings bright golden-orange with fine black stripes and dashes and small white irregular spottings surrounded by black circles (Fig. 2C); each rectangular marking encircled by heavy black border; dorsum of foot cream-white with hundreds of fine black parallel lines radiating from body midline; entire foot edged with bright yellow border; siphon pure white edged with black; ten- tacles ochre-yellow with fine black stripe along entire dorsal side; eyes black; posterior mantle flap ruffled, pure white, bordered with black; penis elongate, smooth, transparent white with prominent, easily seen vas def- erens. Type-locality.—Two m depth off Parcel das Paredes, Abrolhos Reef Com- plex, Bahia State, Brazil (17°47’'S, 39°1'W). Distribution.—At present, known only from the Abrolhos Reefs. 516 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 2. A, Radular row of Acmaea (Collisella) abrolhosensis; B, Living animal of Cyphoma macumba (drawn from life); C, Detail of mantle color pattern of Cyphoma macumba. Stippled area is bright golden orange in life; D, Detail of a juvenile specimen of Murexiella iemanja; length 2 mm; E, Spire of Oliva (Plicoliva) zelindae, showing the well-developed plications. Ecology.—The one living specimen was found crawling on the oak leaf- shaped gorgonian Phyllogorgia dilatata at a depth of 2 m. Etymology.—For the African-based fetishistic Macumba religion that, like the new species in this paper, is indigenous to the coast of Brazil. Remarks.—Interestingly enough, Cyphoma macumba was probably first reported living on Phyllogorgia by Hartt (1870:196). However, he confused it with Cyphoma gibbosum (Linnaeus, 1758), a species not found in the Abrolhos area. The new species is the only known Atlantic Cyphoma with VOLUME 92, NUMBER 3 517 a prominent hornlike projection on the outer lip. In this respect C. macumba closely resembles the Panamic species C. emarginatum (Sowerby, 1830) (Cate, 1973:69, figs. 154—154a). Family Muricidae Genus Dermomurex Monterosato, 1890 Subgenus Trialatella Berry, 1964 Dermomurex (Trialatella) oxum, new species Fig. 1, E and F Material Examined.—HOLOTYPE—length 13 mm, width 7 mm, 25 m depth, 2 km east of Santa Barbara Is., Abrolhos Archipelago, Bahia State, Brazil (17°57'S, 38°41'W), 27 July 1977. USNM 780648. PARATYPES — length 8 mm, USNM 780649; fragment length 5 mm, UMML 8170; both from same locality, depth, and date as holotype. Shell Description.—Stocky, broadly fusiform, with 5 whorls; aperture large, roughly % length of shell; inside of outer lip with 6 elongate teeth that extend well into aperture; siphonal canal short, open, and dorsally recurved; early body whorls with 6 rounded varices; last body whorl with 3 expanded, crenulate, blade-like varices; body whorl sculpture consisting of 3 strong cords with raised pustules and 4 weaker spiral cords; intervarical region with 3 large, rounded knobs, each corresponding to one spiral cord; intri- ticalx white, composed of parallel rows of raised half-cylinders; interior of aperture yellow; operculum unknown. Type-locality.—Twenty-five meters depth, 2 kilometers east of Santa Bar- bara Is., Abrolhos Archipelago, Bahia State, Brazil (19°57'S, 38°41’W). Distribution.—Known only from the Abrolhos Archipelago. Ecology.—The shells were found dead on top of the calcareous mud at the base of the ‘‘chapeirao’’ reef and most probably had rolled down the steep reef face after death. The species apparently lives cryptically in cracks and crevices in the reef. Etymology.—Named for Oxum, principal sea god of the Macumba reli- gion. Remarks.—The only Atlantic Dermomurex that resembles D. oxum is D. abyssicola (Crosse, 1865) from deep water off Guadeloupe, Lesser Antilles. Actually, the new species most closely resembles D. cunninghamae (Berry, 1864) from 40 m depth off Guaymas, Sonora, Mexico. D. oxum can be considered the Atlantic cognate of D. cunninghamae. Genus Muricopsis Bucquoy and Dautzenberg, 1882 Muricopsis oxossi, new species Fig. 1, Gand H Material Examined.—HOLOTYPE—length 9 mm, width 5 mm, 25 m depth, 2 km east of Santa Barbara Is., Abrolhos Archipelago, Bahia State, 518 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Brazil (17°57'S, 38°41’W), 27 July 1977. USNM 780650. PARATYPE— length 7 mm, USNM 780651, from same locality, depth, and date as holo- type. | ) Shell Description.—Small, fusiform, spire high, acute; suture indistinct; sculpture consisting of 5 raised axial cords and numerous close-packed, overlapping scales, giving shell lamellose appearance; varices flattened, winglike, heavily scaled, with 5 flattened spines along edges; spine on shoul- der largest, recurved posteriorly; spines on last varices fused into single large varical wing with serrate edge; 7 varies on last whorl of holotype; color creamy-yellow with 3 slightly darker bands, one at shoulder, one at mid-body, and one at base of siphonal canal; shell covered with irregular white intriticalx; siphonal canal long, fully “4 of shell length; aperture white; inner edge of outer lip with 5 low, rounded teeth; operculum unknown. Type-locality.—Twenty-five meters depth, 2 kilometers east of Santa Bar- bara Is., Abrolhos Archipelago, Bahia State, Brazil (17°57’S, 38°41’ W). Distribution.—Known only from the Abrolhos Archipelago. Ecology.—Like that of the previous muricid, Dermomurex oxum. Etymology.—Named for Oxossi, a sea god of the Macumba religion. Remarks.—Muricopsis oxossi resembles no other known Western Atlan- tic Muricopsis. The new species shows some relationship to M. tulensis Radwin and D’ Attilio, 1976 (Radwin and D Attilio, 1976:233-234), known only from the southern tip of Baja California. Genus Murexiella Clench and Farfante, 1945 Murexiella iemanja, new species Fig. 1, I and J Material Examined.—HOLOTYPE—length 8 mm, width 6 mm, 25 m depth, 2 km east of Santa Barbara Is., Abrolhos Archipelago, Brazil (17° 57'S, 38°41'W), 27 July 1977. USNM 780652. PARATYPE: length 10 mm, USNM 780653, from same locality, depth, and date as holotype. Shell Description.—Stoutly fusiform; spire high, consisting of 2 nuclear and 3 post-nuclear whorls; suture indistinct; body whorl slightly globose; 7 varices per whorl, 5 large foliaceous spines on each varix; spine on shoulder well-developed, twice as long as other varical spines, posteriorly recurved; tips of all spines, except large shoulder spine, distinctly bifurcate; body whorl with 5 raised foliaceous cords corresponding to varical spines; si- phonal canal long, straight, dorsally recurved at tip, with 3 foliaceous cords and short varical spines; aperture ovate; columellar lip adherent posteriorly, detached anteriorly; subsutural region of spire smooth, well-developed, giv- ing shell wide-shouldered, angular appearance; color pinkish-white with dis- tinctive large brown patches between each varix in subsutural region; oper- culum unknown. VOLUME 92, NUMBER 3 319 Type-locality.—Twenty-five meters depth, 2 kilometers east of Santa Bar- bara Is., Abrolhos Archipelago, Bahia State, Brazil (17°57'S, 38 °41’W). Distribution. —Known only from the Abrolhos Archipelago. Ecology.—As in Dermomurex oxum and Muricopsis oxossi—the 2 spec- imens being collected dead in calcareous mud at the base ef a ‘‘chapeirao’’ tec Etymology.—Named for Iemanja, principal sea goddess of the Macumba religion. Remarks.—Murexiella iemanja somewhat resembles M. macgintyi (M. Smith, 1938) from Florida and the Bahamas and M. facetus E. Vokes, 1970, from Aruba, but differs from both by having bifurcate spines, a higher spire, and by its smaller size. Fig. 2D shows a juvenile specimen of M. iemanja, 2 mm in length, composed of the protoconch and one whorl. Family Fasciolariidae Genus Latirus Montfort, 1810 Subgenus Polygona Schumacher, 1817 Latirus (Polygona) ogum, new species Fig. 3, A and B Material Examined.—HOLOTYPE—length 40 mm, width 18 mm, fring- ing reef around Coroa Vermelha, Abrolhos Reef Complex, Bahia State, Brazil (17°57'S, 39°13’W), 29 July 1977. USNM 780654. Shell Description.—Holotype with 7 whorls, each with 8 elongate, low, rounded axial ribs; spiral sculpture on body whorl consisting of 15 raised cords, all equally prominent; siphonal canal long for genus, roughly 4 length of shell; siphonal canal with 3 large, prominent cords, several secondary cords between; columella with 4 folds; inside of lip with 12 rows of large pustules, fusing into lirae deep inside aperture; color orange-red, darker on axial ribs; periostracum thin, reddish-brown, operculum unknown. Type-locality.—In tide pool, west side of fringing reef around Coroa Ver- melha, Abrolhos Reef Complex, Bahia State, Brazil (17°57'S, 39°13'’W). Distribution.—Known only from the Abrolhos Archipelago, but possibly lives elsewhere along the south Bahia and Espiritu Santo coasts. Ecology.—Although the unique holotype was collected dead with a hermit crab, living Latirus ogum probably occurred on the same reef. The fringing reef around Coroa Vermelha contained many large tide pools filled with Shell gravel and algae, an ideal habitat for Latirus. Etymology.—Named for Ogum, a Macumba god often associated with the sea. Remarks.—The ‘‘Latirus sp.’ of Rios (1975: pl. 29, fig. 439) from ‘‘Couves Is.’’ may be a juvenile of L. ogum. Latirus bernadensis Bullock, 1974 (Pl. 2, Fig. C) from the Lesser Antilles and northern Brazil (Bullock, 520 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 3. A, Latirus (Polygona) ogum, dorsal aspect of holotype; B, Latirus (Polygona) ogum, ventral aspect of holotype; C, Latirus bernadensis, holotype; D, Oliva (Plicoliva) ze- lindae, dorsal aspect of holotype; E, Oliva (Plicoliva) zelindae, ventral aspect of holotype. 1974:76—77, fig. 13) is a similar species, but differs from L. ogum in lacking the 3 prominent cords on the siphonal canal, and by having stronger spiral cords, lighter color, and a much more angled shoulder. Latirus ogum very closely resembles the Panamic L. sanguineus (Wood, 1828) (Keen, 1971:615, fig. 1333)—especially so with its red-orange color—and can be considered the Atlantic analogue. VOLUME 92, NUMBER 3 521 Family Olividae Genus Oliva Bruguiere, 1789 Subgenus Plicoliva, new subgenus Diagnosis.—Shell olivid in form, polished, short, rotund, with prominent mamillate protoconch; body whorl with pronounced evenly-spaced, longi- tudinal plications running entire length of shell from spire suture to siphonal canal; plications especially prominent on spire and shoulder (Fig. 2E); col- umellar plications large, 6-12 in number, restricted to lower half of colu- mella. Type-species.—Oliva (Plicoliva) zelindae, new species. Oliva (Plicoliva) zelindae new species Fig. 3, D and E; Fig. 4, A and B Material Examined.—HOLOTYPE—length 31 mm, width 24 mm, Gua- ratibas Reefs, Abrolhos Reef Complex, Bahia State, Brazil (17°25’S, 39°8’W), 19 July 1977. USNM 780655. PARATYPES—length 32 mm, USNM 780656; length 27 mm, my collection; both from same locality and data as holotype; length 26 mm, Museu Nacional do Brasil 3734, Santa Barbara Is., Abrolhos Archipelago (17°58’S, 38°42’W). Shell Description.—Form and sculpture as for subgenus; color pattern composed of network of fine ‘‘tent’’ markings with series of dark, evenly- spaced vertical bars around middle, in turn connected by thin, continuous dark band; spire marked with intermittent dark checkers; color pattern vary- ing from orange-brown to black on white background. Type-locality.—In shell gravel 1 m depth in tide pool, south side of Gua- ratibas Reefs, north end of Abrolhos Reef Complex, Bahia State, Brazil (17°25'S, 39°8’W). Distribution.—Known only from the Abrolhos Archipelago and Reef Complex. Ecology.—The new species appears to live in sand and shell gravel areas in shallow water on and around the larger reefs and islands. Costellaria kaicherae n. sp., Closia largillieri (Kiener, 1841), small Lucina spp. and Corbula spp. were the only mollusks commonly found with O. zelindae on the Guaratibas Reefs. Etymology.—Named for Zelinda Margarida Leao, marine geologist of the Universities of Miami and Bahia, Salvador, Brazil, whose invaluable help made it possible for me to visit the Abrolhos Archipelago in 1977. Remarks.—Oliva (Plicoliva) zelindae is probably the most aberrant of the known Olividae. Having both the longitudinal plications on the body whorl and the enlarged columellar plications, the new species resembles volutes in the genera Lyria or Enaeta more than typical Oliva or Olivella. At first glance, O. zelindae might be taken for a volutid, but the typical olive shape, PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 4. A, Oliva (Plicoliva) zelindae, dorsal aspect of paratype; B, Oliva (Plicoliva) zelindae, ventral aspect of paratype; C, Vexillum (Costellaria) kaicherae, dorsal aspect of holotype; D, Vexillum (Costellaria) kaicherae, ventral aspect of holotype; E, Vexillum (Costellaria) lixa, dorsal aspect of holotype; F, Vexillum (Costellaria) lixa, ventral aspect of holotype; G, Conus iansa, dorsal aspect of holotype; H, Conus iansa, ventral aspect of holotype. high gloss, and multiple columellar plications point to placement in the Olividae. There are no known Caribbean fossil olives that show any relationship to Plicoliva. It appears that the subgenus has been endemic to the Abrolhos area since at least the Miocene. VOLUME 92, NUMBER 3 523 Family Costellariidae Genus Vexillum Roding, 1798 Subgenus Costellaria Swainson, 1840 Vexillum (Costellaria) kaicherae, new species Fig. 4, C and D Material Examined.—HOLOTYPE—length 8 mm, width 3 mm, 1 m depth, tide pool on Guaratibas Reefs, Abrolhos Reef Complex, Bahia State, Brazil (17°25’S, 39°8’W), 19 July 1977. USNM 780657. PARATYPES: lengths 9 mm, 6 mm, USNM 780658; length 9 mm; UMML 8171, fragment length 6 mm, UMML 8172; lengths 9 mm, 9 mm; DMNH 121796; lengths 9 mm, 7 mm, Museu Nacional do Brasil 3735; length 8 mm, collection of Sally D. Kaicher; all from same locality, depth, and date as holotype. Shell Description.—Small, fusiform, elongate, biconic; shoulder rounded; suture well-developed, indented; most adult shells with 5 whorls; proto- conch mamillate, composed of one and one-half whorls; sculpture composed of 15-18 axial ribs, each studded with regularly-spaced round nodules; col- umella with 4 folds; anterior half of shell dark reddish-brown; spire cream- white checkered with occasional brown spots; interior of aperture brown. Type-locality.—One meter depth in tide pool, south end of Guaratibas Reefs, Abrolhos Reef Complex, Bahia State, Brazil (17°25’S, 39°8’W). Distribution.—Known only from the Abrolhos Reef Complex. Ecology.—All specimens were collected dead between sand ripples in tide pools. Other dead shells that accumulated with the new species were Oliva zelindae n. sp., Oliva circinata Marrat in Sowerby, 1871, Conus iansa n. sp., Closia largillieri (Kiener, 1841), and small Lucina and Corbula spp. It is presumed, judging from the fresh condition of the type-specimens, that V. (Costellaria) kaicherae lived in the same tide pools—probably buried in the sand during the day. Etymology.—For Sally Diana Kaicher, malacologist and professional pho- tographer, of St. Petersburg, Florida, in recognition of her work in the family Costellariidae and for her excellent scientific photography, which have both been of great assistance to many scientists. Remarks.—Although the shape and sculpturing of V. (Costellaria) kai- cherae resembles that of typical costellariids, the one and one-half-whorl pro- toconch points to arelationship with the pusiids. Is it possible that V. kaicher- ae and the following new species represent a new subgenus intermediate be- tween the subgenera Costellaria and Pusia. Vexillum (Costellaria) lixa, new species Fig. 4, E and F Material Examined.—HOLOTYPE—length 12 mm, width 5 mm, | m depth in tide pool, west side of Lixa Reef, Abrolhos Reef Complex, Bahia 524 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON State, Brazil (17°43’S, 38°59’W), 23 July 1977. USNM 780659. PARA- TYPES—lengths 19 mm, 12 mm, USNM 780660, both from same locality, depth, and date as holotype. Shell Description.—Fusiform, elongate, biconic; shoulder sharply angled; sculpture with 1S—17 axial ribs per whorl and 17-20 spiral ridges giving shell nodulose appearance; adult shell with 6 whorls; protoconch mamillate, com- posed of one and one-half whorls; columella with 4 folds; spire and shoulder pure white, anterior half of shell rose-pink; interior of aperture pink. Type-locality.—One meter depth in tide pool, west side of Lixa Reef, Abrolhos Reef Complex, Bahia State, Brazil (17°43’S, 38°59’ W). Distribution.—Known only from Abrolhos Reef Complex. Ecology.—Like V. (Costellaria) kaicherae, V. lixa was collected dead between sand ripples in large tide pools. Etymology.—For Lixa Reef (Pedra Lixa), the type-locality—which is named after the lixa, a Bahiano term for any kind of small reef shark. Remarks.—V. (Costellaria) lixa bears an amazing resemblance V. mo- destum (Reeve, 1845) from the Indo-Pacific. The 2 species are very close in shell color, shape, size, and sculpture. However, this similarity must be considered as convergence on similar shell morphologies and does not re- flect phylogenetic or zoogeographic relationships. Although almost identical morphologically, the 2 species have evolved from different stocks: V. mo- destum has a typical costellariid protoconch and arose from the classic Indo- Pacific Costellaria line while V. lixa has a pusiid-type protoconch and, along with V. kaicherae, probably represents a mid-Tertiary off-shoot of Pusia lineage in the South Atlantic. Family Conidae Genus Conus Linnaeus, 1758 Conus iansa, new species Fig. 4, G and H Material Examined.—HOLOTYPE—length 12 mm, width 7 mm, in cal- careous mud, 25 m depth, 2 km east of Santa Barbara Is., Abrolhos Archi- pelago (17°57’S, 38°41’W), Bahia State, Brazil, 27 July 1977, USNM 780661. PARATYPES—length 13 mm, in shell gravel, 1 m depth in tide pool, Lixa Reef, Abrolhos Reef Complex (17°57’S, 39°41’W), 23 July 1977, USNM 780662; length 15 mm, in sand, | m depth off Guaratibas Reefs, Abrolhos Reef Complex (17°25'S, 39°8’W), 19 July 1977, UMML 8173; length 14 mm, same locality, depth, and date as holotype, my collection. Shell Description.—Small, shiny, squat, with wide, heavily coronated shoulder; anterior half of last whorl with 8-15 deeply impressed spiral sulci, posterior half smooth; spire elevated with mamillate protoconch; color vari- able, ranging from white to shades of pink and orange; color pattern com- posed of series of dots and dashes in close-packed spiral rows overlaid with VOLUME 92, NUMBER 3 525 large amorphous patches of darker color; some specimens greyish-white with bright white color pattern; spire with alternating patches of darker color; interior of aperture white; periostracum thin, smooth, transparent yellow, with small shaggy tufts on shoulder. Animal.—Foot and body translucent grey with numerous small white patches; proboscis tipped with black; eyes black; operculum tiny, oval. Type-locality.—Twenty-five meters depth, 2 kilometers east of Santa Bar- bara Is., Abrolhos Archipelago, Bahia State, Brazil (17°57'S, 38°41'W). Distribution.—The new cone appears to be restricted to the Abrolhos Archipelago and Reef Complex. Ecology.—Conus iansa prefers a soft substrate, such as calcareous mud or sandy gravel, on or near the reef complex. It most probably feeds on small polychaete worms. Associated mollusks include: Turritella exoleta (Linnaeus, 1758), Turbinella laevigata Anton, 1839, Oliva (Plicoliva) zelin- dae n. sp., Oliva circinata Marrat in Sowerby, 1871, Nassarius capillaris (Watson, 1882), and Mitrella albovittata Lopes, Coelho, and Cardoso, 1965. Etymology.—Named for Iansa, a Macumba goddess often associated with both the moon and the sea. Remarks.—Conus iansa resembles no other known Western Atlantic cone Shell. In some aspects, such as shell coloring and the coronated spire, the new species resembles some forms of C. otohimeae Kuroda and Ito, 1961, from Japan and Taiwan. Conus iansa may be allied to the Conus magel- lanicus Hwass, 1972—Conus speciosissimus Reeve, 1848, species complex of the West Indies; this assumption is based on the small shell size and strongly coronated shoulder. Members of this complex are always associ- ated with hard-bottom communities on Caribbean reefs, and the soft-bottom habitat of C. iansa is unusual. Besides C. iansa, only 2 other species of cone shells are known from the Abrolhos region; C. brasiliensis Clench, 1942 (Van Mol, Tursch, and Kempf, 1967:237—238, map 2) and C. jaspideus Gmelin, 1791. Acknowledgments I would like to thank Mr. Arnaldo C. dos Santos Coelho, Museu Nacional do Brasil, Rio de Janeiro, for loan of study material from the Abrolhos Archipelago, and Mrs. Zelinda Margarida Leao, Universities of Bahia, Bra- zil, and Miami, for allowing me the opportunity of joining her on her doctoral research trip to the Abrolhos. Special thanks are given to Sally Diana Kaich- er, St. Petersburg, Florida, for the excellent photographs used in this paper, to Dr. Gilbert L. Voss, Rosenstiel School of Marine and Atmospheric Sci- ence, University of Miami, for critical review and encouragement in my research, and to Dr. Donald R. Moore, also of the Rosenstiel School, for help in sorting of material. 526 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Literature Cited Bullock, Robert C. 1974. A contribution to the systematics of some West Indian Latirus (Gastropoda: Fasciolariidae).—The Nautilus 88(3):76—77, fig. 13. Cate, Crawford Neill. 1973. A systematic revision of the recent Cypraeid Family Ovulidae (Mollusca: Gastropoda).—The Veliger 15 (Supplement): 69, figs. 154—154a. Hartt, Charles Frederick. 1870. Geology and physical geography of Brazil. Chapter IV. The islands and coral reefs of the Abrolhos.—Thayer Expedition Report. Fields, Osgood and Co., Boston, pp. 174-214. Keen, A. Myra. 1971. Sea shells of tropical West America.—Second Edition. Stanford Uni- versity Press, Stanford, California. p. 615, fig. 1333. Laborel, Jacques. 1969. Les Peuplements De Madreporaires Des Cotes De L’Etat De Bahia Et Les Abrolhos.—Annales De L’ Université D’ Abidjan, Série E, II, Fascicule 3, Ecol- ogie:125, 131, 135-142, figs. 45, 58, 50-55. Radwin, George E. and Anthony D’Attilio. 1976. Murex shells of the World. An illustrated guide to the Muricidae.—Stanford University Press, Stanford, California. pp. 233-234. Righi, G. 1966. On the Brazilian species in the Acmaea subrugosa complex.—Malacologia 4(2):269-295, plates 1-2, figs. 1-2, 12-14. Rios, E. C. 1975. Brazilian Marine Mollusks Iconography.—Fundagao Universidade do Rio Grande, Museu Oceanografico. p. 104, figs. 437, 439. Smith, E. A. 1890. Notes on the zoology of Fernando de Noronha.—Lin., Soc. Jour. Zool. 10:479-S03, pl. 30. Van Mol, J-J., B. Tursch and M. Kempf. 1967. Mollusques Prosobranches: Les Conidae du Bresil. Résultats Scientifiques des Campagnes de la ‘‘Calypso.”’ Annal. l’ Inst. Oceanogr. XLV(16):237—238, map.2. Rosenstiel School of Marine and Atmospheric Science, University of Miami, 4600 Rickenbacker Causeway, Miami, Florida 33149. PROC. BIOL. SOC. WASH. 92(3), 1979, pp. 527-532 VOORTHUYSENIELLA VENTRESSI, NEW SPECIES (MICROPROBLEMATICA) FROM THE BASAL PLEISTOCENE OF LOUISIANA Drew Haman Abstract.—A new species of Voorthuyseniella Szczechura, V. ventressi, (1969), is described from subsurface Pleistocene strata on the Louisiana continental shelf. Observations are made on the morphologic and paleo- bathymetric significance of the taxon. A form of Voorthuyseniella Szczechura, 1969, with a distinctly com- pressed test has been described, in open nomenclature, from the subsurface Pliocene of the Texas continental shelf (Haman, 1977). The recovery of 2 additional specimens of the taxon in 2 wellbore samples from early Pleis- tocene strata on the Louisiana continental shelf (West Delta area) provides additional information which allows a more definitive taxonomic diagnosis that provides taxonomic stability for this Plio-Pleistocene organism. Other occurrences of Voorthuyseniella in the Neogene-Recent strati- graphic interval of the Gulf of Mexico province have been documented by Keij, 1970 (offshore Texas, Recent); Haman and Kohl, 1976 (offshore Tex- as, Pleistocene); Haman, 1978 (offshore Texas and Louisiana, Pleistocene); Haman, in press (offshore Louisiana mudlumps, late Quaternary). The standardized test orientation, morphologic and measurement termi- nology are based on Haman (1978:190). Voorthuyseniella ventressi, new species Fig. 1 Description.—Test calcareous, unilocular, as high as wide, tapering from base of camera to porta, camera cone-shaped in side view (Fig. 1b,g). Cam- era extends laterally to either side of a median line between the lateral apertures. Camera ovate when viewed from above (Fig. 1d) and below (Fig. le). The ovate, compressed camera is situated transverse to the tubus. Cam- era is widest (0.37 mm) just below the middle. Tubus moderately flat, slightly raised in the direction of lateral aperture ‘A’ (Fig. 1b). Tubus slightly more narrow at the middle, in the area of the tubus pore, compared to the ends at the lateral apertures (Fig. le). A subparallel-sided furrow is present on the tubus and extends from the tubus pore to either side ending just beyond the camera limits (Fig. le). Porta subcircular to elliptical in shape (0.035 mm xX 0.025 mm) situated in a slightly bevelled depression and surrounded 528 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON by an elliptical abraded area (0.09 mm x 0.07 mm) (Fig. 1d). Porta orien- tated transverse to the lateral apertures and in line with the greatest camera width. Porta is eccentric, not located at camera top, but is slightly offset and directed towards lateral aperture ‘A’ (Fig. 1b). Tubus pore subcircular (0.020 mm X 0.015 mm) situated between the lateral apertures and in line with them. Tubus pore does not extend beyond the tubus furrow width (Fig. le). Lateral aperture ‘A’ subcircular to ovate (0.045 mm x 0.035 mm) (Fig. la) with a poorly defined flap-like structure over it (Fig. Id). A rim (0.018 mm in width) surrounds the aperture. Lateral aperture “B’ subcircular, 0.045 mm diameter (Fig. 1c) surrounded by a rim 0.015 mm in width. No septa are evident. Test surface appears striate and punctate under light micro- scope examination but under SEM examination the striae are shown to result from differential camera solution, and punctae are elongate depres- sions on test wall, not true punctae. The shallow elongate depressions are arranged concentrically around the porta and are restricted to the camera. Solution effects are evident over both camera and tubus but appear to be more severe on tubus. Test dimensions.—Maximum length 0.47 mm, maximum width 0.37 mm, height 0.36 mm. Locality.—Well cutting sample, West Delta area, offshore Louisiana. Stratigraphic level.—Early Pleistocene, Zone N.22 of Blow (1969). Repository.—In author’s collection pending final disposition. Etymology.—The new species is named in honor of W.P.S. Ventress, Senior Staff Paleontologist, Chevron U.S.A., New Orleans, in recognition of his many years of providing guidance to numerous paleontologists in Gulf Coast biostratigraphy. Remarks.—The specimens ascribed to this species (holotype, paratype, plus the form previously described in open nomenclature) exhibit variation in their morphology, which is believed to be the result of specific variation _ and possibly related to the growth process and/or solution effects. The camera ranges in shape from a narrow elongate cone (Fig. 1b ) to a shorter, more robust type (Fig. 1g). The form previously described (Haman, 1977) appears to be an intermediate type. Cameral compression ranges from se- vere (Fig. Id, e) to more moderate (Fig. li, j). Again the form previously detailed appears to be intermediate. The porta ranges in shape from dis- tinctly elongate (Haman, 1977, Pl. 1, fig. 3; Pl. 2, fig. 3) to less elongate = Fig. 1. Voorthuyseniella ventressi. a-e, Holotype; f-j, Paratype: a, f, Lateral aperture A’; b,g, Side view (lateral aperture ‘A’ at left); c, h, Lateral aperture ‘B’; d, i, View to porta and supra apertural ‘flap’ (lateral aperture ‘A’ at left); e, j, View to tubus, tubus pore and tubus furrow (lateral aperture ‘A’ at left). Bar = 100 um. VOLUME 92, NUMBER 3 530 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON (Fig. 1d) to subcircular (Fig. 1i). The porta shape and size in these specimens appear directly related to the degree of test dissolution around the feature. It appears that the shallow pits described earlier undergo selective disso- lution with a collapse type of feature being subsequently developed around the porta, thus changing the shape and size of the porta, as suggested by Haman (1977, Pl. 2, fig. 3). The holotype and paratype exhibit different degrees of solution. The holotype is well preserved with little evidence of differential solution, while the reverse is the case with the paratype. The shape of the small tubus pore may also be affected by a similar phenomenon, giving rise to slight shape and size variation. The size ranges of various morphologic features are noted below: Haman 1977 Specimen Holotype Paratype Pl. 1, Figs. 1-3, Fig. la—3 Fig. 1f-g Pl. 2, Figs. 1-3 (entire) (partly broken) (broken) Max. length 0.47 mm 0.47 mm 0.46 mm Max. height 0.36 mm 0.40 mm 0.43 mm Max. width 0.37 mm 0.38 mm 0.44 mm Porta 0.04 x 0.03 mm 0.50 x 0.03 mm 0.06 x 0.03 mm The specimens described herein exhibit no definite affinity to any previously described forms of Voorthuyseniella. It should be noted that due to the fragmented nature of the test, lateral apertures ‘A’ and ‘B’ were reversed in the description of the form previously detailed (Haman, 1977). All the previously documented Pleistocene-Recent occurrences of Voor- thuyseniella in the Gulf of Mexico province listed earlier, comprised forms of this taxon that possess a globular or elongate-globular (hemispherical) test. This type of morphology is typically associated with a shallow water (<100 fathoms) type of environment (see Keij, 1970; Hantzschel, 1975). One occurrence of a globular form in paleobathymetrically deeper waters (>100 fathoms) was attributed to downslope displacement of the organism (Ha- man, 1978:190). The Voorthuyseniella specimen described from the Texas Pliocene was suggested to have developed such a morphology (i.e. flattened camera) in response to a paleobathymetric parameter (Haman, 1977:135). The philos- ophy of this hypothesis has been detailed (ibid) and need not be repeated here. The foraminiferid assemblage associated with that occurrence un- questionably indicated a paleobathymetry of 250-500 fathoms (Haman, 1977:135). The Pleistocene forms described herein do not, by means of foraminiferid association, attain middle bathyal depths as did the Pliocene form, but a paleobathymetry in at least the upper bathyal range is indicated. The fora- VOLUME 92, NUMBER 3 531 miniferid assemblage in the sample containing the holotype was composed of 22 benthic species with in excess of 130 individuals, and 8 planktic species with over 100 individuals. The sample containing the paratype possessed a richer foraminiferid assemblage composed of 38 benthic and 9 planktic species, both groups with an excess of 300 individuals. These assemblages by analogy to the data of Phleger (1951); Parker (1954); Murray (1973); LeRoy and Levinson (1974); and LeRoy and Hodgkinson (1975) indicate an upper bathyal depth range (>100-—<250 fathoms). The assemblage from the latter sample suggests an environment in the lower reaches of the upper bathyal range. It is believed that these data support the hypothesis that the forms of Voorthuyseniella with a compressed test are more indicative of deeper bathymetries than the more globular test types of this genus. Acknowledgments I am indebted to Dr. W. H. Akers and Mr. W. P. S. Ventress for reading an early draft of this manuscript; to Mr. B. Kohl for relevant discussions; to Mr. D. Greig for expert assistance with the SEM photography; to Mr. J. R. Bailey and Ms. E. Spencer for technical assistance; to Chevron U.S.A., Inc. (Eastern Region) for allowing access to the SEM; and to Chevron Oil Field Research Company for publication permission, and defrayment of publication charges. Literature Cited Blow, W. H. 1969. Late middle Eocene to Recent planktonic foraminiferal biostratigraphy.— In P. Bronnimann and H. H. Renz (eds.), Proceedings of the First International Con- ference on Planktonic Microfossils, Geneva, 1:199-—421. Haman, D. 1977. Pliocene Voorthuyseniella (Problematica) from the Texas Continental Shelf. —Tulane Stud. Geol. Paleont. 13(3):135-140. . 1978. Voorthuyseniella Szczechura 1969, (Problematica), microfossile du Pléistocene du plateau continental de Louisiane et du Texas.—Rev. Micropaléont. 20(4): 187-196. . In press. Late Quaternary Microproblematica (Voorthuyseniella) from a Mississippi Delta mudlump.—Tulane Stud. Geol. Paleont. , and B. Kohl. 1976. Voorthuyseniella (Problematica) from the Pleistocene of the Texas Continental Shelf.—Tulane Stud. Geol. Paleont. 12(3):157—160. Hantzschel, W. 1975. Treatise on Invertebrate Paleontology, Part W, Miscellanea, Supple- ment 1, Trace Fossils and Problematica, 2nd ed. (ed. C. Teichert), Microproblematica.— Geol. Soc. America and Univ. Kansas, W153—W168. Keij, A. J. 1970. Taxonomy and stratigraphic distribution of Voorthuyseniella (Problematica) I and IJ.—Koninkl. Nederl. Akad. Weten. Proc., Ser. B, 73:479-499. LeRoy, D. O., and K. A. Hodgkinson. 1975. Benthonic Foraminifera and some Pteropoda from a deep-water dredge sample, northern Gulf of Mexico. —Micropaleontology 21(4):420-447. , and S. A. Levinson. 1974. A deep water Pleistocene microfossil assemblage from a well in the northern Gulf of Mexico.—Micropaleontology 20(1):1-—37. Murray, J. W. 1973. Distribution and ecology of living benthic foraminiferids.—Heinemann Educational Books, xiii + 274 pp. Doz PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Parker, F. L. 1954. Distribution of the Foraminifera in the northeastern Gulf of Mexico.— Harvard Mus. Comp. Zool. Bull. 111(10):453-588. Phleger, F. B. 1951. Ecology of Foraminifera, north-west Gulf of Mexico. Pt. 1, Foraminifera distribution.—Geol. Soc. America Mem. 46: 1-88. Szczechura, J. 1969. Problematic microfossils from the upper Eocene of Poland.—Rev. Espan. Micropaleontologica, 1(1):81—94. Chevron Oil Field Research Company, P.O. Box 446, La Habra, Califor- nia 90631. PROC. BIOL. SOC. WASH. 92(3), 1979, pp. 533-550 A REVISION OF THE MILLIPED GENUS DELOPHON, WITH THE PROPOSAL OF TWO NEW TRIBES IN THE SUBFAMILY ABACIONINAE (CALLIPODIDA: CASPIOPETALIDAE) Rowland M. Shelley Abstract.—The callipodid milliped genus Delophon is revised. Three species are recognized: D. georgianum Chamberlin, the type species, D. serrulatum Causey, and D. holti, n. sp., known only from Hamblen Co., Tennessee. Delophon carolinum Hoffman is relegated to the synonymy of the type species. The three species are entirely allopatric and range from the Blue Ridge Province of western North Carolina and the Cumberland Mountains of Tennessee to the Gulf Coastal Plain of Alabama. A disjunct population of D. georgianum also occurs in the Kings Mountain region of North Carolina, which is some 50 miles east of the Blue Ridge Front in the Piedmont Plateau. Delophon georgianum and D. serrulatum are anatomi- cally very similar and appear to be sister species; D. holti, however, appears to represent another evolutionary line. Generically, Delophon is related to Abacion and Tetracion, the other genera of the subfamily Abacioninae, but gonopodal differences warrant segregation into a different tribe. Keys to the tribes and genera of the Abacioninae and species of Delophon are presented, along with gonopod illustrations and a range map. The milliped order Callipodida is well represented in the epigean fauna of the southeastern United States by species of the distinctive genus Abacion. These large, brown-black, crested diplopods are commonly encountered in a variety of native and urban habitats in all physiographic provinces. Oc- curring sympatrically with Abacion in Tennessee, Alabama, and the south- ern Blue Ridge Province is a second callipodid genus, Delophon, which is smaller in size but displays the same general body coloration. Adults of Delophon closely resemble juveniles of Abacion, but can be distinguished by the absence of legless segments posteriorly, the more translucent exo- skeleton, and by the crest and gonopodal characters set forth in the ensuing key. Individuals of Delophon are also much stiffer and harder to unroll without breaking than the juveniles of Abacion with which they can be confused. The two genera also appear to differ ecologically, as I have always found Delophon in moist litter, often near streams, and never in the dry pine and urban environments that Abacion frequents. The third southeast- ern callipodid genus, Tetracion, is exclusively troglobitic. 534 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Unlike many North American milliped genera, the history of Delophon is short and simple. It was erected by Chamberlin (1943) in his typical fashion, with only three sentences of diagnosis, for a species from Gaines- ville, Georgia, which he named georgianum. Chamberlin did, however, note the principal diagnostic feature, the enlargement of the coxal process of the male gonopod into a sheath around the “‘telopodite,’’ thus allowing for fu- ture recognition of the genus. Hoffman (1950) proposed a second name, D. carolinum, for specimens found at Highlands, North Carolina, and Causey (1954) described an additional species, D. serrulatum, from the Mobile Bay region of Alabama. The only other treatment is the listing of Chamberlin and Hoffman (1958). Hoffman (1979) assigned Abacion, Tetracion, and De- lophon to the subfamily Abacioninae in the family Caspiopetalidae and re- ported that Delophon consisted of three species ranging from North Caro- lina to Alabama. The purpose of this paper is to describe the genus Delophon, update the nomenclature of the established species and summarize their distribution, and diagnose a new species from Hamblen Co., Tennessee. The occasion is also taken to propose two new tribal categories for the Abacioninae. I am grateful to Richard L. Hoffman for the privilege of describing the new species and for comments on an early draft of the manuscript, and to him and William A. Shear for loan of material in their collections. The holotype of D. serrulatum Causey was kindly loaned by Norman I. Platnick, Amer- ican Museum of Natural History (AMNH), and that of D. carolinum Hoff- man, by Ralph E. Crabill, Jr., United States National Museum (USNM). Access to the type of D. georgianum, in the Chamberlin collection (RVQ), was also courtesy of Dr. Hoffman. I also thank Drs. Platnick, Herbert W. Levi, Museum of Comparative Zoology, Harvard University (MCZ), and Howard V. Weems, Jr., Florida State Collection of Arthropods (FSCA), for access to specimens in their care. Material in the North Carolina State Museum collection is indicated by the acronym NCSM. This study was supported in part by financial aid from the National Science Foundation, Grant. No. DEB-7702596. KEY TO TRIBES AND GENERA OF ABACIONINAE 1. Pore crests small and narrow, subequal in size to primary crests, not distinctly enlarged anteriad; male gonopods without flagella; coxal processes thin and translucent, subequal in length to and sheathing PARP RRP VOmera GS basdt pada Ih eee en Ge ena Delophonini, new tribe Sia gaa sg ao he ca MSA sass SEALE Sch vo te: Sun MIS Mere CNT Delophon Chamberlin — Pore crests considerably larger than primary crests, distinctly en- larged anteriad; male gonopods with flagella; coxal processes rela- VOLUME 92, NUMBER 3 535 tively thick and opaque, shorter than and subparallel to femora ie rere A metres “ieee. Fav the Slt of mise! oh, Abacionini, new tribe 2. Dorsum with only four primary crests between pore crests on all segments; flagellum of male gonopods relatively short, not extending to distal elements of telopodite; troglobitic species with elongated legs and antennae, reduced pigmentation, and fewer ocelli ........ AOE MiB ELON AIR i doe Sa Fires Perse Ded wrecave eal 34s Tetracion Hoffman — Dorsum with four primary crests between pore crests through seg- ment 11, six primary crests thereafter; flagellum of male gonopods relatively long, extending to level of distal elements of telopodite; large pigmented epigean species, with normal legs, antennae, and WECM AIA sept shaadi elds Bes Bien jeectt yew iamyy ard co aes, de Abacion Rafinesque Abacionini, new tribe Components.—Abacion Rafinesque, 1820; Tetracion Hoffman, 1956. Diagnosis.—A tribe of moderate-large callipodids with the following char- acteristics: pore crests distinctly larger than primary crests, ozopores lo- cated in widened anterior portion of crests; male gonopods possessing fla- gella, coxal processes shorter than and subparallel to femora. Range.—Eastern United States east of central Great Plains, from Illinois and eastern Nebraska south to central Texas and east to New York and peninsular Florida. Species occur in subterranean and epigean environ- ments. Abacion occurs in a wide variety of surface habitats in the eastern United States and is one of the most common indigenous, urban diplopods. Delophonini, new tribe Component.—Delophon Chamberlin, 1943. Diagnosis.—A tribe of small callipodids with the following characteris- tics: pore crests subequal in size to primary crests, only slightly widened anteriad, ozopores located in latter portion of crests; male gonopods without flagella, coxal processes thin and translucent, subequal in length to and sheathing telopodite femora. Range.—Southern United States, from Cumberland Mountains of central Tennessee and Blue Ridge Mountains of western North Carolina to Gulf Coast of Alabama. Species occur exclusively in epigean habitats. Genus Delophon Chamberlin Delophon Chamberlin, 1943:13. Causey, 1954:63. Chamberlin and Hoffman, 1958: W112) 536 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Type species.—Delophon georgianum Chamberlin, 1943, by original des- ignation. Description.—A genus of small callipodids with the following character- istics: body composed of head and 47-51 segments at maturity. Head broad- ly rounded between o¢ellaria; suture visible only in epicranial region; frons and clypeus flattened, subtrapezoidal, narrowing towards labrum, width across genae less than that across epicranium; ocellaria variably subtrian- gular, with prominent Organ of Tomosvary on inner margin just above an- tennal socket; antennae relatively long and slender, composed of seven segments, 3 and 6 the longest, with four sensory cones apically. Body segments cylindrical in cross section, slightly flattened dorsad; all segments with numerous longitudinal ridges or crests beginning on caudal half of collum, medial crests on collum slightly shorter than lateral crests; segments with 10 dorsal crests between pore crests through segment 11, 14 dorsal crests thereafter; all crests running entire length of metaterga, some (primary) more distinct than others (secondary). Ozopores beginning on segment 6, located anteriorly in a dorsolateral crest; pore crests slightly larger than primary crests, forming a narrow V with point caudad. Dorsal setal formula as follows (method of Hoffman and Lohmander [1964]): Segment Setal Series med. a b © de lat: 1 AAAAA D AAAAA 3 AAAAA 4 AAAAA 5 Ar CPP Pp SAPP 6 Po PL PWETE Legs with 7 segments; coxae of legs 3—4 of males and 2-10 of females with prominent ventral pads; all legs with prominent, curved claw. Hypo- proct tripartite, with 2 prominent setae on quadratic median lobe and 1 spinneret projecting from under distal edge of each lateral lobe. Paraprocts divided by oblique suture into smaller posterior and larger anterior sclerites. Gonopods (terminology of Hoffman [1954, 1956]) proportionally large, clearly visible externally, femur projecting caudad from segment 7 and post- femur reflexed abruptly laterad or anteriolaterad; with a large, laminate tibiotarsus and shorter, narrower parasolenomerite; postfemoral process present or absent; solenomerite located between and variously shielded by tibiotarsus and parasolenomerite, apically blunt or acute; course of prostatic groove distinctly demarcated by black pigmentation; coxal process thin and translucent, greatly enlarged and expanded into broad sheath partially sur- rounding femur, subequal in length to femur; flagellum completely absent; coxae connected by prominent sclerotized sternum (Fig. 4). VOLUME 92, NUMBER 3 537 Distribution.—Southern United States from the Blue Ridge Province of western North Carolina south to the Gulf Coastal Plain of Alabama and west to the Cumberland Mountains of Tennessee. Species.—Three are known, but it seems possible that an additional species may be discovered in central Tennessee-Kentucky. Causey (1954) presented a key to the callipodid genera in the Gulf Coastal states and to the species of Delophon recognized at that time. The following couplets are based on more definitive characters and incorporate nomenclatorial changes made herein. KEY TO SPECIES OF DELOPHON (BASED ON ADULT MALES) 1. Distal elements of gonopods forming angle of approximately 45° with femur; parasolenomerite parallel to and only slightly shorter than tibiotarsus; solenomerite short and blunt, concealed by parasolen- omerite and tibiotarsus; coxal process (sheath) with distomedial edge jagged, slightly expanded laterad apically ....................... 2 — Distal elements of gonopods forming angle of about 90° with femur; parasolenomerite much shorter than and not parallel to tibiotarsus; solenomerite much larger than parasolenomerite, broad basally and spiniform apically, clearly visible between parasolenomerite and tibiotarsus; coxal process (sheath) with distomedial edge smooth, strongly expanded laterad behind tibiotarsus ............ holti, n. sp. ZL OSUCMOrAl,DTOCESS PRESEN .........5. 0-608 georgianum Chamberlin =F EOStremOral PrOCESS ADSENE 0. Fok ec os caine oer serrulatum Causey Delophon georgianum Chamberlin Figs. 1-3 Delophon georgianum Chamberlin, 1943:13, figs. 28-30. Chamberlin and Hoffman, 1958:112. Delophon carolinum Hoffman, 1950:18, pl. 5, fig. 4. Chamberlin and Hoff- man, 1958:112. Wray, 1967:153. NEW SYNONYMY. Type specimen.—Male holotype (RVC) collected by Wilton Ivie, 24 April 1943, from Gainesville, Hall Co., Georgia. Diagnosis.—Distinguished by following features of male gonopods: ele- ments distal to femur forming approximately 45° angle with femur; coxal process with distomedial edge jagged, distolateral margin entire, sheathing femur; postfemoral process present, spiniform, about 4 length of tibiotar- sus, acuminate; tibiotarsus long and acuminate, cupped at midlength to ac- commodate solenomerite; solenomerite short, blunt, located between par- asolenomerite and tibiotarsus. 538 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON VOLUME 92, NUMBER 3 539 Holotype.—The following details are supplemental to the original descrip- tion by Chamberlin (1943). Color of preserved specimen brown, fading laterally at poriferous carinae, epicranium and interantennal region dark mottled brown, fading out in clyp- eal region. Middorsal stripe absent from collum, beginning on protergum of segment 2. Relative lengths of antennomeres as follows: 2=3 >5=4>6>7> 1, 2-6 clavate. Collum with about 20 crests of uniform height beginning near midlength, anterior half of collum smooth. Gonopods (Figs. 1-3) with broadly curved femur projecting caudad from segment 7 to prozonite of segment 9; postfemur bent abruptly anterolaterad, expanding into broad tibiotarsus and parasolenomerite, extending to edge of segment 7; coxal process sheathing femoral region, extending caudad to level of apical femoral bend, distomedial margin jagged. Postfemoral process spiniform, approximately 4 length of tibiotarsus, extending to just beyond level of separation of tibiotarsus and parasolenomerite and running parallel to latter processes, closely appressed to postfemoral suface. Tibiotarsus a large, acute, folded structure, cupped at midlength to partially sheath so- lenomerite. Parasolenomerite relatively straight, acute, slightly shorter than and parallel to tibiotarsus, expanded slightly at level of solenomerite; latter a short, blunt, bisinuately curved structure, flared apically with translucent subapical spur, located between parasolenomerite and tibiotarsus at ap- proximately midlength of both these structures. Variation.—The postfemoral process varies in length being extremely short on a male from Madison Co., NC (NCSM A359) and directed obliquely on a male from Wayah Bald, Macon Co., NC (RLH), so that it is almost perpendicular to the axis of the postfemur and tibiotarsus. Ecology.—As noted in the introduction, this species is usually found in moist leaf litter, often near streams. Although common in the southern Blue Ridge Province, only a few individuals are usually found at one time. The largest samples I have seen contain only seven specimens. Distribution.—With one notable exception, the range of D. georgianum coincides closely with the boundaries of the southern Blue Ridge Province. All the material from Tennessee has come from the eastern extremity of the state near the North Carolina border; none has been taken from the Ridge and Valley Province only a few miles to the west. The northernmost au- <_— Figs. 1-3. D. georgianum, left gonopod of holotype. 1, medial view. 2, caudal view. 3, lateral view. Scale line = 0.5 mm. co, coxal process (sheath); f, femur; pfp, postfemoral pro- cess; pS, parasolenomerite; s, solenomerite; tt, tibiotarsus. 540 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON thentic record (with male specimens) is from the edge of the mountains in Wilkes Co., NC, and the southernmost is from the type locality, which is near the mountains but actually in the Piedmont Plateau of north Georgia. The most striking aspect of the distribution of D. georgianum is the occur- rence of a disjunct population some S50 miles east of the Blue Ridge Front in the Kings Mountain region of North Carolina. This is a small area of mountains in the Piedmont Plateau containing peaks of over 1,000 ft. and is one of five inselberg areas in the state (Filka, 1978). No specimens have been collected in the intervening lowlands between the Kings Mountain region and the Blue Ridge Mountains, and the Kings Mountain population appears to be truly isolated from the remainder of the species. I can detect no anatomical differences, however, and find no reason to recognize this population taxonomically. This type of disjunct distribution in the southeast, in which isolated populations of predominantly montane species occur in the Piedmont, is obtained in a number of plant species, most notably the hemlock (Tsuga canadensis) and the white pine (Pinus strobus). Hardin and Cooper (1967) concluded that these plant disjuncts are relicts of a continuous Pleistocene forest which became dissected as climates warmed during post- glacial time. Perhaps the Kings Mountain population of D. georgianum is likewise a relict from a time when the species was more widely distributed and has since disappeared from most of its range due to climatic changes. Survival in the Kings Mountain region could have resulted from a slightly cooler microclimate afforded by the peaks and coves in this area. Specimens have been examined as follows: Georgia: Habersham Co., Clarksville, M, F, 31 July 1958, R. L. Hoffman (RLH). Hall Co., Gainesville, M holotype, 24 April 1943, W. Ivie, (RVC) TYPE LOCALITY. White Co., along Tray Mtn. Rd. near McClure Cr., M, 24 June 1975, D. A. Rossman (AMNH). Rabun Co., N side Rabun Bald, near road to Rabun tower, M, F, 30 May 1964, H. R. Steeves (FSCA); and Glade Mtn., F, 27 July 1949, R. L. Hoffman (RLH). Tennessee: Monroe Co., 16 mi. SE Madisonville, campground on Citico Creek road, Cherokee National Forest, M, juv. F, 12 October 1978, R. M. Shelley and W. B. Jones (NCSM A2471). Sevier Co., Elkmont, Great Smoky Mountains National Park (GSMNP), 2 M, 2 F, 16-22 September 1959, W. Suter (FSCA); 10.1 mi. S Gatlinburg, 3,800’ near small forest stream, GSMNP, 2 M, 24 May 1977, A. Newton, and M. Thayer (MCZ); => Figs. 4-6. D. serrulatum. 4, ventral view of gonopods of male specimen from Walker Co., GA, showing sternum (st). 5-6, left gonopod of specimen from Sequatchie Co., TN. 5, lateral view. 6, caudal view. Abbreviations as in Figs. 1-3. Scale line = 0.36 mm for Fig. 4, 0.43 mm for Figs. 5-6. 542 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 4.6 mi. S Gatlinburg 2,300’, mixed hardwood forest, GSMNP, 2 F, 24 May 1977, A. Newton and M. Thayer (MCZ); Chimneys Picnic Area, 7.3 mi. S Gatlinburg, GSMNP, M, 16 May 1978, R. M. Shelley and W. B. Jones (NCSM A1892) and 5 F, 28 July 1949, R. L. Hoffman (RLH); Newfound Gap, 5,000’ GSMNP, juv. M, 25 May 1962, L. Hubricht (RLH). North Carolina: Macon Co., Coweeta Hydrological Station near Otto, 4 F, 1 August 1958, R. L. Hoffman (RLH), 2 M, 2 F, 27 September 1964, H. R. Steeves (FSCA), and M, 6 F, 23 May 1965, W. Suter (FSCA); Highlands, M, F, 21 July 1949, R. L. Hoffman (RLH); Buck Cr. clearcut, 1.6 mi. NW intersection Buck Cr. Rd. and co. rd. 1538, M, 2 July 1976, F. A. Coyle (NCSM A2129); and Wayah Bald, M, F, 3 July 1954, R. L. Hoffman (RLH). Jackson Co., Cullowhee Gap, M, 3 F, 10 July 1958, R. L. Hoffman (RLH); farm near Webster, M, 16 May 1970, W. A. Shear (WAS); and Cherokee Indian Reservation near Blue Ridge Parkway, F, 3 October 1960, W. J. Gertsch and W. Ivie (AMNH). Graham Co., Joyce Kilmer Memorial Forest, M, 3 F, 25 September 1971, F. A. Coyle (NCSM A2089); 4.8 mi. SSE Robbinsville, along co. rd. 1127, 0.2 mi. W jct. co. rd. 1116, F, 25 July 1974, R. M. Shelley (NCSM 2465); 2.8 mi. WSW Robbinsville, along co. rd. 1115, 2.5 mi. S jet. co. rd. 1120, M, F, 25 July 1975, R. M. Shelley (NCSM 2745). Swain Co., Deep Creek, 4 mi. N Bryson City, M, F, 1 October 1960, W. J. Gertsch and W. Ivie (AMNH); base Cliff Ridge near Nantahala, M, 6 May 1951, L. Hubricht (RLH); and Smokemont, GSMNP, F, 15 May 1978, R. M. Shelley and W. B. Jones (NCSM A1881). Haywood Co., GSMNP, Cataloochee Area, F, 8 July 1976, R. M. Shelley (NCSM A919). Transyl- vania Co., Thompson River Gorge SE Lake Toxaway, F, 5 September 1961, R. L. Hoffman (RLH); and Pink Beds Recreation Area, 8 mi. N Brevard, M, 5 F, 28 July 1958, R. L. Hoffman (RLH). Buncombe Co., N Bat Cave, 3 F, 5 July 1955, R. L. Hoffman (RLH). Madison Co., along US hwy. 25- 70, 0.9 mi. N Hot Springs, M, 22 July 1975, R. M. Shelley and J. C. Clamp (NCSM A359); Paint Rock, M, 13 September 1952, L. Hubricht (RLH); and along US hwy. 70, 0.8 mi. NE jct. NC hwy. 22, F, 23 July 1971 (RLH). Mitchell Co., below Carver’s Gap on Roan Mtn., 5,000’, 2 F, 23 September 1950, L. Hubricht (RLH); and Altapass, M, 3 F, 20 May 1957, R. L. Hoff- man (RLH). Wilkes Co., 7.2 mi. NW North Wilkesboro, along co. rd. 1541, 0.1 mi. E jet. co. rd. 1544, M, 12 August 1975, R. M. Shelley and J. C. Clamp (NCSM A466). Gaston Co., Crowders Mtn., M, 3 March 1950, D. L. Wray (FSCA); 4.8 mi. SW Gastonia, along co. rd. 1131, 0.2 mi. W. ject. co. rd. 1133, M, 29 April 1976, R. M. Shelley (NCSM A701); 4.4 mi. W Gastonia, along co. rd. 1122, 1.1 mi. E jct. cord. 1131, M, 10 July 1976, M. Filka and W. W. Thomson (NCSM A1123); 2.5 mi. S Bessemer City, along co. rd. 1125, 0.1 mi. S jet. US hwy. 74-29, M, F, 17 October 1976, M. Filka and G. Wicker (NCSM A2249) and M, 2 F, 9 April 1977, M. Filka (NCSM A2170); W of Gastonia on co. rd. 1106, 1.5 mi. E jet. co. rd. 1236, M, 16 VOLUME 92, NUMBER 3 543 October 1976, M. Filka and G. Wicker (NCSM A2312); and 7 mi. SE Bes- semer City, along co. rd. 1103, jct. co. rd. 1112, M, 2 F, 9 April 1977, M. Filka (NCSM A2316). Delophon serrulatum Causey Figs. 4-6 Delophon serrulatum Causey, 1954:64, figs. 1-3. Chamberlin and Hoffman, 1958:112. Type specimen.—Male holotype (AMNH) collected by N. B. Causey, 5 January 1954, from an unknown locality near Loxley, Baldwin Co., Ala- bama. The label in the vial states ‘‘Loxley, Baldwin Co., Mississippi,’ but Causey (1954) reported that the locality was 9 miles west of Loxley at the junction of US hwy. 90 and Miss. state hwy. 89. Chamberlin and Hoffman (1958) corrected the state to Alabama but restated that the site was nine miles west of Loxley. The road map for Baldwin Co., however, does not show a state highway 89, and even if there were one, it would be east of Loxley rather than west, since the state highways numbered in the 80’s are all east of the town. In view of this confusion, it seems best to consider the type locality as being Loxley vicinity, Baldwin Co., Alabama. Diagnosis.—Male gonopods similar in general appearance to those of D. georgianum except postfemoral process absent. Holotype.—The following details are supplemental to the more complete description by Causey (1954). Color of preserved specimen generally pale brown, fading laterally; epi- cranium a light mottled brown, interantennal region dark brown, fading out in clypeal region. Middorsal stripe absent from collum, beginning on pro- tergum of segment 2, expanding slightly on segments 4—6, widest on seg- ments 12-30, narrowing gradually thereafter to penultimate segment. Relative lengths of antennomeres as follows: 2=3 >5>4>6>7> 1, 2-6 clavate. Ocellaria roughly triangular, about 34 vocelli in irregular rows. Anterior half of collum smooth, crests beginning near midlength. Setae on collum located either on smooth portion or anterior portions of crests, none at posterior position. Gonopods (Figs. 4—6) similar in size and configuration to those of D. georgianum, except postfemoral process entirely absent; subapical spur of solenomerite with edge deeply serrate. Distribution.—Delophon serrulatum ranges from the southern Gulf Coast- al Plain to the Cumberland Plateau of northern Alabama to the Ridge and Valley Province of Tennessee. It is completely allopatric with D. georgian- um. Specimens have been examined as follows: Alabama: Baldwin Co., Loxley vic., jct. AL hwy. 89 and US hwy. 90, M 544 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON holotype, 5 January 1954, N. B. Causey and D. Causey (AMNH) TYPE LOCALITY. Coneucuh Co., 3 mi. W Brooklyn, outside Turk Cave, M, 11 August 1965, S. B. Peck (FSCA). Talladega Co., 6 mi. SW Sylacauga, M, 18 September 1959, W. Suter (FSCA). Shelby Co., Oak Mountain State Park, 2 M, 3 F, 16 October 1960, H. R. Steeves (FSCA), and 2 M, 3 F, 16 April 1965, N. B. Causey (FSCA). Marion Co., Hackleburg, 2 M, 5 F, 25 May 1964, J. W. King (FSCA). Winston Co., near Natural Bridge Cave, M, 8 June 1965, collector unknown (FSCA). Morgan Co., 3.5 mi. SE Fayette- ville, M, F, 21 May 1972, S. B. Peck (WAS). Madison Co., Woodville, 2 M, 3 F, 7 September 1965, S. B. Peck (FSCA). Franklin Co., the Dismals, M, F, 17 June 1961, H. R. Steeves (FSCA), and M, 4 September 1961, W. Suter (FSCA). Jackson Co., Princeton, M, 29 October 1960, H. R. Steeves (FSCA); and 5 mi. NE Garth, 7 M, 6 F, 19 May 1972, S. B. Peck (WAS). Georgia: Dade Co., Cloudland Canyon State Park, M, 15 April 1962, H. R. Steeves (FSCA), and M, F, 17 June 1962, H. R. Steeves (FSCA). Walker Co., 5 mi. SE Lafayette, M, 10 June 1967, S. B. Peck (WAS). Tennessee: Marion Co., 2.2 mi. SE Monteagle, M, 9 May 1954, L. Hu- bricht (RLH). Sequatchie Co., 5 mi. SE Dunlap, wet woods in limestone ravine, M, 20 August 1956, R. L. Hoffman (RLH). Fentress Co., James- town, Jordan Motel, 3 M, 3 F, 16 June 1962, H. R. Steeves (FSCA). Remarks.—The remarkable similarity in structure between D. serrulatum and georgianum suggests that they may be only geographic races of a single species. However, no specimens have been collected in the intervening area, and there is no alternative but to recognize two valid species until such material is obtained. My personal feeling is that the two forms are indeed reproductively isolated and that their present status is correct. Delophon holti, new species! Figs. 7-9 Type specimens.—Male holotype and two male and one female paratypes (RLH) collected by R. L. Hoffman and P. C. Holt, 1 June 1952, from moist, wooded sinkholes +1 mi. SE Russellville, Hamblen Co., Tennessee. One male paratype deposited in NCSM collection. Diagnosis.—Distinguished by following features of male gonopods: ele- ments distal to femur forming approximately a right angle with femur; coxal process with distomedial edge smooth, sheathing femur and extending lat- erad about 14 length of tibiotarsus; postfemoral process absent; parasolen- omerite comparatively small, much shorter than and not parallel to tibi- 1 This species was first recognized as new by Richard L. Hoffman, who generously allowed me to describe it, requesting only that it be named for Perry C. Holt, professor emeritus of biology at Virginia Polytechnic Institute and State University, Blacksburg, Virginia. VOLUME 92, NUMBER 3 545 aN 8 Figs. 7-9. D. holti, left gonopod of holotype. 7, caudal view. 8, lateral view. 9, distal portion of coxal process (sheath), ventral view. Abbreviations as in Figs. 1-3. Scale line = 0.5 mm. 546 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON APSR ENON FSO TES POLE ER le RSS pastes Pee Ne ME ea TS nen fe mi: Ses Mas Pela Vr ee peo Ti @ D. georgianun ee ia 2 ea @ D. serruiatum ae ’ @ D. holti J } + / : AN ber i Ann | | ree f aoe Tae Se ae PP es NE ia i Fig. 10. Distribution of Delophon. otarsus, arising from ventral surface of solenomerite; solenomerite large, broad basally and narrowing to acute, spiniform tip; tibiotarsus mostly broad, angling abruptly caudad apically into spiniform projection. Description.—Length about 19 mm, greatest width about 1.1 mm; ca. 47 segments. Preserved specimens very pale in color, crests alone showing brown pigmentation; epicranium light mottled brown, interantennal region darker, fading out in clypeal region. Middorsal stripe absent from collum but present on remaining segments. Antennae long and slender, extending beyond caudal margin of segment 5; relative length of antennomeres 2 = 3>5>4>6>7> 1, 2-6 clavate. About 28 ocelli in subtriangular patch. Collum with 17 crests beginning near midlength, medial crests shorter than lateral ones, anterior half of collum smooth. Secondary crests subequal to primary crests in height and length up to about segment 11, fading out VOLUME 92, NUMBER 3 547 behind this segment; transition from 10-14 dorsal crests between pore crests on segment 12. Gonopods (Figs. 7—9) small, femur extending caudad only to metazonite of segment 8; postfemur perpendicular to femur; tibiotarsus extending lat- erad to edge of body, distance between tibiotarsal apices subequal to width of body; coxal process very broad, sheathing femoral region, expanded laterad apically for about 1% length of tibiotarsus, acuminate, edges not serrate. Postfemur very short, without postfemoral process. Tibiotarsus di- rected laterad, broadly curved and folded, concave at mid-length to accom- modate solenomerite, angling abruptly caudad apically and narrowing into spiniform projection. Parasolenomerite relatively short, arising from ventral surface of solenomerite, broad basally but narrowing rapidly to acuminate tip, directed caudolaterad, not parallel to tibiotarsus. Solenomerite relative- ly large, broadly expanded basally but narrowing rapidly to acuminate tip, tip located between tibiotarsus and parasolenomerite, directed dorsolaterad. Ecology.—In 1978 I visited the type locality in an unsuccessful attempt to find topotypes. This is a dry, rocky area that has largely been converted into pasture and is poor milliped habitat. The dominant trees are red cedars (Juniperus virginiana). The type specimens were found in a moist sinkhole, suggesting that D. holti may have specialized ecological requirements and occupy a particular niche which has been overlooked by field collectors. Distribution.—Known only from the type locality. Remarks.—As in D. georgianum, there are three terminal processes on the gonopods of D. holti, but this is because the solenomerite of the latter is large and spiniform. The solenomerite of the type species is a short blunt spur off the parasolenomerite and nearly concealed by the latter structure and the tibiotarsus. The postfemoral process of D. georgianum, one of the three terminal branches referred to by Causey (1954), is absent from D. holti. Thus, the three terminal branches of the gonopods of D. holti and D. georgianum are different and not an indication of any phylogenetic close- ness between the two species. Distribution The distribution of each species of Delophon (Fig. 10) has been summa- rized in the appropriate account, and only one additional point needs to be emphasized. The three species are entirely allopatric, with no overlap in their known ranges. Additional collecting in the Cumberland Mountains and Plateau, which have not been intensively sampled, may reveal sympatry of D. holti and serrulatum, but I feel confident that only the type species occurs in the Blue Ridge Province. The thorough collecting in this area by myself and others would surely have produced another species by now if 548 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON holti georgianum Serrulatum Fig. 11. Relationships in the genus Delophon. one occurred there. Additional specimens of D. holti are needed to provide increased knowledge of its range, gonopodal variation, and the general bi- ology of the species. It is possible, however, that the situation with Delo- phon may be analogous to that of the spirostreptid genus Choctella, in which there is one common, widely dispersed species, C. cumminsi Cham- berlin, and one with a very restricted range, C. hubrichti Hoffman (Hoff- man, 1965). It is worth noting in this regard that D. holti has only been collected once, some 27 years ago, a fact which suggests both a narrow range and reduced abundance. Relationships Generic: Relationships among North American callipodid genera are un- clear. This topic was not discussed in the most definitive account (Loomis, 1937), at which time Delophon was unknown. Hoffman (1954) provided the foundation for such a consideration in his proposed terminology of calli- podid gonopods, and he later (1956) mentioned that the male genitalia of Tetracion indicated affinity with both Abacion and Delophon. It seems clear that the affinities of Delophon are with the other two eastern genera and not with Colactis, Heptium, and the other western callipodids, which seem to represent a separate evolutionary line. The eastern genera all have a well developed tibiotarsal blade which projects laterad at a distinct angle from the femur, as opposed to being an expansion along the axis of the femur. VOLUME 92, NUMBER 3 549 In Tetracion and D. georgianum and serrulatum, the angle between the femur and tibiotarsus is approximately 45°, whereas in Abacion and D. holti, it is about 90°. Delophon is unique, however, in the development of the coxal process into a large, translucent sheath around the femur and in the absence of the flagellum, which has been variously called a coxal process (Hoffman, 1954), sternal process (STP) (Hoffman, 1956), and coxosternal process (Shear, 1974). Specific: The many gonopodal similarities—most notably a roughly 45° angle between the distal elements and the femora; the short, blunt solen- omerites; and the long, blade-like parasolenomerites and tibiotarsi—point to a close affinity between D. georgianum and serrulatum. Their only dif- ference involves the postfemoral process (present in D. georgianum, absent in D. serrulatum), and they seem to be sister species, only one step removed from a common ancestor. Delophon holti, however, is so entirely different from its congeners that it must represent a second line from some earlier source. That D. holti exists suggests the possibility of at least one additional undiscovered species of Delophon, its sister species, perhaps in central Tennessee-Kentucky. Investigators conducting field studies in this area should be alert to this possibility. These hypothesized relationships in the genus Delophon are depicted in Fig. 11. Literature Cited Causey, Nell B. 1954. Three new species and new records of southern millipeds.—Tulane Stud. Zool., 2(4):63—68. Chamberlin, Ralph V. 1943. On some genera and species of American millipeds.—Bull. Univ. Utah Biol. Ser., 8(2): 1-20. , and Richard L. Hoffman. 1958. Checklist of the millipeds of North America.—Bull. U.S. Natl. Mus. No. 212, 236 pp. Filka, Marianne E. 1978. Studies on the Diplopoda of the Kings Mountain region.—Unpub- lished Masters Thesis, Zoology Department, North Carolina State University, Raleigh, North Carolina, 114 pp. Hardin, James W., and Arthur W. Cooper. 1967. Mountain disjuncts in the eastern Piedmont of North Carolina.—J. Elisha Mitchell Sci. Soc., 83:139-S0. Hoffman, Richard L. 1950. Records and descriptions of diplopods from the southern Appa- lachians.—J. Elisha Mitchell Sci. Soc., 66:11-33. . 1954. A new milliped of the genus Colactis from Mexico (Chordeumida: Lysiopetal- idae).—Am. Mus. Nov. No. 1673, 4 pp. 1956. New genera and species of cavernicolous diplopods from Alabama.—Geol. Surv. Alabama, Mus. Pap. No. 35, 11 pp. 1965. A second species in the Diplopod genus Choctella (Spirostreptida: Choctelli- dae).—Proc. Biol. Soc. Wash., 78:55—58. . 1979. Classification of the Diplopoda.—Museum d Histoire Naturelle, Geneva, Swit- zerland. , and Hans Lohmander. 1964. The Diplopoda of Turkey, parts I and II.—Mitt. Hamburg Zool. Mus. Inst., 62:101-151. 550 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Loomis, Harold F. 1937. Crested millipeds of the family Lysiopetalidae in North America, with descriptions of new genera and species.—Proc. U. S. Natl. Mus., 84:97-135. Shear, William A. 1974. North American cave millipeds. II. An unusual new species (Dory- petalidae) from southern California, and new records of Speodesmus tuganbius (Tri- chopolydesmidae) from New Mexico.—Calif. Acad. Sci., Occ. Pap. No. 112, 9 pp. Wray, David L. 1967. Insects of North Carolina, Third Supplement.—North Carolina De- partment of Agriculture, Division of Entomology, Raleigh, 181 pp. North Carolina State Museum, P.O. Box 27647, Raleigh, North Carolina 27611. PROC. BIOL. SOC. WASH. 92(3), 1979, pp. 551-571 A SYNOPSIS OF THE MILLIPED GENUS CAMBALA, WITH A DESCRIPTION OF C. MINOR BOLLMAN (SPIROSTREPTIDA: CAMBALIDAE) Rowland M. Shelley Abstract.—The spirostreptid milliped genus Cambala is summarized; an- nulata (Say), minor Bollman, texana Loomis, ochra Chamberlin, speobia (Chamberlin), washingtonensis Causey, and hubrichti Hoffman are recog- nized as valid species. The remaining six nominal species are relegated to appropriate synonymies. Cambala minor is described and illustrated for the first time, and the posterior gonopods of all species except washingtonensis are compared, to establish their identities. The latter is known only from females, and its identity is still obscure. The occurrence of Cambala in the Pacific northwest is confirmed, as authentic specimens have been collected in southwestern Oregon, and the range of annulata is extended to the vi- cinity of Pittsburgh, Pennsylvania. Cambala is widespread and common in unglaciated portions of the eastern United States from southwestern Penn- sylvania and southern Hlinois to the Gulf Coastal Plain, ranging westward to the Edwards Plateau region of west Texas and adjoining areas. The milliped genus Cambala is the only representative of the family Cam- balidae in the eastern United States. In the southern Appalachian Mountains and southern Atlantic States, individuals of C. annulata (Say) and C. hu- brichti Hoffman are a dominant element of the litter fauna, being more prevalent in the cooler months of spring and fall. The distinctions between these two large species were detailed by Hoffman (1958), who also clarified their nomenclature. The situation is quite different in the Cumberland Pla- teau and midwestern states, however. Small to moderate sized cambaloids range southwestward into Texas and adjacent areas and are common in the surface and cave faunas, but their identities are uncertain and their nomen- clature, confusing. Miscellaneous records exist in a number of papers, but none can be considered reliable. The only comprehensive treatment is that of Loomis (1938), but several additional species have been described, and there have been other nomenclatorial changes. Today, Cambala is com- prised of the following 12 species, one of which is divided into two subspe- cies. I here recognize only seven of these as valid, the others placed in synonymy as indicated. In the ensuing discussions they will be referred to only by the specific or specific and subspecific names. S52 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON annulata (Say, 1821) minor Bollman, 1888 arkansana Chamberlin, 1942 cara Causey, 1953 texana Loomis, 1938 ochra Chamberlin, 1942 saltillona Chamberlin, 1943 loomisi (Hoffman, 1956) speobia (Chamberlin, 1952) reddelli reddelli Causey, 1964 reddelli inornatus Causey, 1964 washingtonensis Causey, 1954 hubrichti Hoffman, 1958 The major difficulty in the taxonomy of Cambala has been uncertainty about the identity of minor, which was originally described as a subspecies of annulata by Bollman (1888a). He published no illustrations, as he never did in any of his papers, and reported no diagnostic characters. His only comment was that the new form was ‘‘similar to C. annulata, but much smaller and of a yellowish-brown shade.’’ Loomis (1938) missed a golden opportunity to resolve the problem when, in elevating minor to a full species, he neglected to examine the type specimens and based his conclu- sion entirely on the meaningless drawing of Williams and Hefner (1928) for a form from Ohio. Loomis’ action was correct, though risky without viewing the type material, but he did not clarify the identity of the species. All subsequent species of Cambala have likewise been described without any- one’s examing Bollman’s types, which have rested all these years at the U.S. National Museum. Consequently, synonymies have developed, and the confusion is such that the only species which can be accurately identified today are annulata and hubrichti due to Hoffman’s treatment (1958). My interest in this problem developed after receiving several shipments of midwestern cambaloids for identification. After examining the types of Cambala annulata minor, it was immediately obvious that cara, for which an illustration of the posterior gonopod had been published (Causey, 1953), was a synonym. Since the type locality of arkansana is only a few counties east of that of cara, it seemed likely that this species too was identical to minor. The question thus arose as to how many more synonymies existed among the 12 nominal species of Cambala. Causey (1964) disposed of caeca Loomis and captiosa Causey by bringing them under speobia, but it seemed that the genus could be reduced still further. Consequently, I have under- taken a review of Cambala to clarify the nomenclature and provide, for the first time, a definitive description of minor. Types of all species including caeca and captiosa have been examined except that of washingtonensis, VOLUME 92, NUMBER 3 553 which is absent from its published repository, the American Museum of Natural History (AMNH). Chamberlin and Hoffman (1958) suggested that this species was probably not congeneric with the others, but I have seen crested cambaloids from Oregon at the AMNH that closely resemble the eastern species externally and key out to Cambala in Loomis’ key (1938). They resemble speobia in the configuration of the posterior gonopods, and suggest that washingtonensis may be a valid species of Cambala after all. The results of this study indicate that there are only six unquestionably valid species of Cambala; washingtonensis is here treated as valid but may prove to be a synonym of speobia. Since washingtonensis was based on a female, topotypical males are needed to resolve its status. Synonymies, type localities, and known ranges are listed herein for each species of Cambala, along with miscellaneous comments and illustrations of the posterior gon- opods. The configuration of the posterior gonopod, particularly the coxal process, seems the only reliable character for identifications. Overall size is of some value, and there are three distinct classes, with minor and ochra being intermediate between annulata and hubrichti, the largest species, and speobia and texana, the smallest. Cambala annulata and hubrichti can be separated by somatic features (Hoffman, 1958), but I have been unable to discern external differences between the other species. The species reported herein are those that have been placed in Cambala by past authors. No new species were discovered among the hundreds of specimens examined, and it seems unlikely that any more exist east of the Mississippi River. I have not attempted to deal here with the larger issue of distinctions between the various Nearctic cambaloid genera or redefinition of the genus Cambala. Shear (1969) suggested that both Mexicambala and Troglocambala should be regarded as synonyms of Cambala, but Causey (1971) asserted that the former was distinct based upon ‘‘well defined so- matic and sexual characters.’’ Shear later (1973) agreed with this assess- ment. I have evaluated Troglocambala and concur that it is not a valid genus. Some of the distinguishing characters of Troglocambala are insig- nificant, whereas others vary between individuals. The reduced ornamen- tation of the dorsum, for example, is found frequently in Cambala, which suggests that some of the “‘smoother’’ western genera may not be valid. A critical review of all Nearctic cambaloid genera is badly needed and should receive a high priority when future revisions of North American diplopods are considered. Acknowledgments I am grateful to the following curators for loan of type specimens in their care: Ralph E. Crabill, Jr., U.S. National Museum; Norman I. Platnick, American Museum of Natural History; and Eric H. Smith, Field Museum 554 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON of Natural History. Access to the types in the Chamberlin collection was kindly provided by Richard L. Hoffman, who also loaned material in his personal collection and reviewed an early draft of the manuscript. Speci- mens in the Florida State Collection of Arthropods were studied with per- mission of Howard V. Weems, Jr., and those in the Museum of Comparative Zoology, Harvard University, through the courtesy of Herbert W. Levi. I also thank William A. Shear and Charles P. Withrow for loan of cambaloids in their collections. This study was aided by financial assistance from the National Science Foundation, Grant No. DEB-7702596. Acronyms cited in the text to denote the sources of preserved study material of minor are as follows: AMNH—American Museum of Natural History, New York, NY; FSCA—Florida State Collection of Arthropods, Gainesville, FL; MCZ—Museum of Comparative Zoology, Harvard Uni- versity, Cambridge, MS; NCSM—North Carolina State Museum, Raleigh, NC; RLH—Private collection of Richard L. Hoffman, Radford, VA; RVC— Private collection of the late Ralph V. Chamberlin, now being accessioned by the U.S. National Museum; WAS—Private collection of William A. Shear, Hampden-Sydney, VA. Genus Cambala Gray Cambala Gray, 1832:784, pl. 135, figs. 2, 2a—c. Bollman, 1887:38; 1893:120. McNeill, 1888:10. Williams and Hefner, 1928:123. Loomis, 1938:36-—37. Causey, 1953:155; 1959a:234; 1964:237-238; 1971:273. Chamberlin and Hoffman, 1958:173. Eclytus (nec Holmgren, 1855) Chamberlin, 1952a:10-11. Eclomus Chamberlin, 1952b:71. Chamberlin and Hoffman, 1958:175. Troglocambala Hoffman, 1956:9-10. Type-species.—Of Cambala, C. lactaria Gray, 1832 (= Julus annulatus Say, 1821), by monotypy; of Eclomus, E. speobius Chamberlin, 1952b, by original designation; of Troglocambala, T. loomisi Hoffman, 1956, by orig- inal designation. Species.—Seven, distinguishable by the configuration of the anterior cox- al lobes of the posterior male gonopods, as shown in Figs. 5—10. Hoffman (1958) produced a couplet based on somatic and gonopodal characters sep- arating annulata and hubrichti, the two largest species, which are sympatric in the southern Appalachians. However, I have been unable to detect any reliable crest or other somatic characters which can separate all the species of Cambala. Range.—tThe distribution of Cambala has never been precisely defined. Loomis (1938) stated that it was the most widespread of the American cam- baloid genera, occurring in many of the eastern states, Texas, and Arkansas. Causey (1964) described the range as being the United States from western VOLUME 92, NUMBER 3 555 ACL Figs. 1-4. Cambala minor lectotype. 1, left first leg, lateral view. 2, anterior gonopods, cephalic view. 3, left anterior gonopod, lateral view. 4, left posterior gonopod, lateral view. ACL, Anterior Coxal Lobe; CP, Coxal Plate; F, Flagellum; PCL, Posterior Coxal Lobe; S, Sternum; T, Telopodite. Scale line = 0.1 mm for figs. 1, 3, and 4; 0.125 mm for fig. 2. 556 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Washington and northern Idaho to the Atlantic Coast, with a center of abundance in the Appalachian Mountains. As shown in the inset map of Fig. 13, Cambala is widespread in unglaciated portions of the eastern United States from southwestern Pennsylvania-southern Illinois south to the Gulf Coastal Plain and northern Florida. Westward, the range extends through Louisiana and Arkansas to New Mexico and the Edwards Plateau region of west Texas. Its occurrence in the Pacific northwest can now be confirmed, due to the material from Oregon at the AMNH. Cambala annulata (Say) Figs. 5, 11 Julus annulatus Say, 1821:103—104. Bollman, 1893:145. Cambala lactaria Gray, 1832:784, pl. 135, figs. 2, 2a—c. Cambala lactarius Gervais, 1837:48. Cambala annulata Cope, 1869:181. Bollman, 1887:38-39; 1888b:339. McNeill, 1888:10. Chamberlin, 1918:24; 1947:58; 1952c:30. Brimley, 1938:498. Hoffman, 1958:91-92. Wray, 1967:158. Shelley, 1978:54—56, fig. 44. Cambala cristula Loomis, 1938:39, fig. 12; 1939:168; 1943:390. Causey, 1953:156; 1959a:234. Chamberlin and Hoffman, 1958:174. Spirobolus annulatus Wood, 1865:212. Type locality.—Southeastern United States, probably in Atlantic Coastal Plain between Charleston, South Carolina, and Jacksonville, Florida (Hoff- man, 1958). Range (Fig. 11).—The range cited by Hoffman (1958), central Virginia to northern Florida and inland from unglaciated portions of Ohio and Indiana to Alabama, can now be expanded to include southwestern Pennsylvania. Specimens from Pittsburgh, Allegheny Co., are present in the MCZ collec- tion. Several males and females from Bryn Mawr, Montgomery Co., are available at the FSCA, but the complete absence of any specimens from Maryland, District of Columbia, and northern Virginia indicate that this is surely an error. In all the years O. F. Cook was at the USNM he never found one individual in the DC area. Additional material from Bryn Mawr = Figs. 5-10. Left posterior gonopods of six species of Cambala, lateral views, setation omitted. 5, annulata, specimen from Chatham Co., NC. 6, hubrichti, specimen from Cocke Co., TN. 7, minor lectotype, specimen from Bloomington, Monroe Co., IN. 8, ochra, specimen from Hart Co., KY. 9, speobia, specimen from Williamson Co., TX. 10, texana holotype, specimen from Nacogdoches Co., TX. ACL, Anterior Coxal Lobe; PCL, Posterior Coxal Lobe; T, Telopodite. Scale lines apply to the figures on either side and = 0.5 mm for figs. 5-6, and 0.1 mm for figs. 7-8 and 9-10. 558 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON is needed before this record can be given any credence. The map (Fig. 11) encompasses all known locality records including those amassed since Hoff- man’s account (1958). The record of Chamberlin (1952a) from Asheville, Buncombe Co., North Carolina, could be either annulata or hubrichti, as I have collected both species from near this city. Ecology.—Hoffman (1958), reporting on montane specimens of annulata, stated that individuals tended to be rather secretive and occurred in the deepest levels of humus or burrows of small mammals. The most popular habitat was moist areas in dry upland oak woods, but the species was oc- casionally found under partly buried logs and rotting stumps in dry sites. In eastern Piedmont North Carolina, annulata is more prevalent in the cooler weather of spring and fall and is common in deep leaf piles. The milliped is hard to find in the summer, however, and only one juvenile has been col- lected in August (Shelley, 1978). Remarks.—Although I have not conducted an exhaustive study of the gonopods of annulata, they do seem to be quite homogeneous throughout the range. Therefore, it seems that Loomis’ name, cristula, relegated to the synonymy by Hoffman (1958), is destined to remain there and not be res- urrected for a geographic race in the Appalachian region. Loomis (1943) reported an anomalous male from Jackson Co., Florida, which had two sets of apparently normal gonopods in typical apertures on segments 7 and 8. The location of this individual is unknown; I could not find it at the MCZ in August 1978, and this was the repository of most of the material on which his paper was based. Assuming Loomis’ description is correct, this would be an example of heteromorphosis in addition to those I reported in 1977 and the only abnormal non-polydesmid known from North America. Cambala minor Bollman Figs. 1-4, 7, 12 Cambala annulata minor Bollman, 1888a:404—405. Cambala minor Loomis, 1938:40. Chamberlin and Hoffman, 1958:174-175. ?Cambala minor Loomis, 1943:390. Causey, 1959a:234—235; 1963:79. Shear, 1969:135, fig. 1. Cambala annulata (nec Say, 1821) Bollman, 1887:38—39; 1888b:339; 1888c, 1. Williams and Hefner, 1928:123, fig. 17B. ?Cambala annulata (nec Say, 1821) McNeill, 1888:10. Cambala arkansana Chamberlin, 1942:3, figs. 2-3. Chamberlin and Hoff- man, 1958:174. NEW SYNONYMY. Cambala cara Causey, 1953:156, figs. 13-14. Chamberlin and Hoffman, 1958:174. NEW SYNONYMY. Type specimens.—Bollman deposited a type series of two males and elev- en females in the U. S. National Museum without designating a holotype. VOLUME 92, NUMBER 3 559 A male was selected from these syntypes by Dr. Hoffman on 12 January 1957, and I officially designate it the lectotype of Cambala annulata minor. The type series was collected by C. H. Bollman from Bloomington, Monroe Co., Indiana, on an unspecified date, which was probably between 1885 and 1887. Diagnosis.—A moderate sized species of Cambala distinguished by two uncate projections from the anterior coxal lobe of the posterior gonopod and by the distal concavity on the anterior edge of the posterior coxal lobe of this structure. Lectotype.—Body long and slender, slightly narrower in neck region, with approximately 64 segments, segment 4 and last two segments without legs. Specimen tightly coiled, length unmeasurable, width 1.5 mm. Color yellowish as noted by Bollman (1888b), caudal portions of dorsal crests slightly brownish. Head with 6 ocelli in linear row on each side. Relative lengths of anten- Nomeness 2) =) 52226 > Si] 40h 7: Collum smooth and shiny, minutely granular, extending forward to back edge of ocellar row; subequal in length to next two segments. Low dorsal crests beginning on segment 2, becoming progressively larger and reaching maximum height on segment 5, terminating on penultimate segment; 4 dorsal crests between peritremata; approximately 11-12 crests laterally on sides of body, becoming thinner and less distinct ventrally. Peritremata beginning on segment 5 and continuing to antepenultimate segment, pear shaped, pos- terior portion subequal in width and height to dorsal crests, anterior portion about twice as broad and slightly more elevated; pore in center of anterior portion. Surface between crests smooth, minutely granular. Epiproct broadly rounded, not extending beyond valves; paraprocts reen- trant; hypoproct short and very broad, margin straight, lateral corners cov- ered by small processes emerging from beneath terminal segment. First legs (Fig. 1) short, 6 segments, proximal two segments widest, ter- minal claw absent. Anterior gonopods (Figs. 2—3) separated by high, apically divided ster- num, coxal plates curving distad behind telopodites, apically divided and expanded caudad into thin laminae; telopodites apically divided with lateral portion moderately expanded caudad; flagella arising at posterobasal edges of laminae of coxal plates. Posterior gonopods (Figs. 4, 7) with anterior coxal lobes terminating in two short uncate projections arising on medial and lateral sides; posterior lobes with typical hamate setae, blunt, distal concavity on anterior edge; telopodites extending slightly beyond caudal margin of gonopod. Paralectotypes.—Although the gonopods are fully developed, the male paralectotype is not fully grown, with only 50 segments, the last five legless. The dimensions are length 22.1 mm, width 1.3 mm. The most conspicuous 560 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON variation among the type specimens is the size of the crests, which are much sharper and more distinct on the paralectotypes than on the lectotype. The latter has the lowest and most rounded crests of any specimen of C. minor I have seen. Variation.—There is some color variation in the species. After 90 years in alcohol, the type specimens are still yellowish, but other specimens are gray, approximately the same color as annulata. Legs 4—7 were damaged in the dissections of both male type specimens, and the following observations were taken from the male from Greene Co., OH, whose gonopods are identical to those of the lectotype. Legs 4—5 are normal, but the femora of legs 6—7 have small lobes on the ventral sides. The lobe on leg 6 is slightly larger. The anterior process of the posterior gonopod varies with respect to the height of the flattened portion between the two hooked projections. On the lectotype it is low, resulting in two distinct hooked projections. On some specimens, however, it is higher and produced into a blunt tooth anteriad. The two hooked projections always point ventrad, except for a male from Jeff Davis Co., Miss., in which they curve mediad to the base of the process. Ecology.—Cambala minor is a troglophilic milliped which can live and reproduce in epigean and subterranean habitats. In Arkansas Causey (1953) recorded it from several surface habitats including pine forests near creeks, dry oak and oak-hickory hillsides, and under driftwood on a marshy creek bank. Causey (1959a) reported finding it on bat, rat, and raccoon dung, wet gravel, and in humus in the entrance and twilight zones of caves in the Cumberland Plateau, but some of these may have been specimens of ochra instead. Distribution (Fig. 12).—Cambala minor occurs widely in the east-central region of the United States, from the Ridge and Valley Province of western Virginia to the Ozark region of Arkansas, north to south-central (unglaciat- ed) Ohio-Indiana and south to the Gulf Coastal Plain. Specimens have been examined as follows; ‘“‘female only’’ samples are reported from areas where minor is the only small cambaloid known to occur. Virginia: Alleghany Co., McElwee’s Cave, ° Feb. 1950, R. L. Hoffman (RLH). Tazewell Co., Burke’s Garden, 2, 20 April 1957, R. L. Hoffman (RLH). Bath Co., Clark’s Cave, 6, 15 April 1960, J. R. Holsinger (FSCA). Augusta Co., Glade Cave, juv. d, °, 1 April 1960, J. R. Holsinger (FSCA); and Cane Hill Cave, 2 6, Nov. 1961, J. R. Holsinger (FSCA). Washington Co., Fraley’s Cave, Wyndale, ° , 6 December 1960, C. W. Greever (FSCA). West Virginia: Pendleton Co., 1 mi. NE Mouth of Seneca, 2, 30 May 1952, L. Hubricht (RLH). Greenbriar Co., Higgenbotham Cave, ?, 11 April 1957, T. C. Barr and D. F. Black (RLH); Arbuckle School Cave, Max- VOLUME 92, NUMBER 3 561 wellton, 6, 2, S. B. Peck, 26 June 1968 (WAS); Ludington Cave, 2000’ from Bell entrance, 3d, 2 October 1965, J. M. Rutherford (FSCA): and Rapps Cave, 3d, 2, 13 May 1962, J. R. Holsinger (FSCA). McDowell Co., Windmill Gap near McComas, 2, 15 April 1973, W. M. McKenzie (WAS). Kentucky: Carter Co., along U. S. hwy. 460, 5 mi. W Grayson, 2, 22 April 1960, R. L. Hoffman and J. G. Barker (RLH). Bell Co., Pine Mountain State Park, wildflower garden, 3, 4 2, Feb. 1976, J. K. Ettman (NCSM A797). Clark Co., along Boone’s Creek near Athens, 4 6, 4 2, 11-23 Oc- tober 1976, T. C. Barr and J. Reddell (FSCA). Hart Co., Rosebury near Cane Cr., 3 ¢6, 5 October 1957, L. Hubricht (RLH). Barren Co., Short Cave Sink, 2.2 mi. NW Park City, 4 6, 3 2, 27 April 1975, L. Hubricht (RLH). Edmonson Co., Green R. floodplain near Styx R., 9 3, several fragmented 2 2, 20 October 1956, L. Hubricht (RLH). Ohio: Greene Co., Yellow Springs, Glen Helen, ¢, 2, 16 November 1968, A. A. Weaver (WAS). Indiana: Jefferson Co., 3.5 mi. W Madison, 4 6,5 2, 10 March 1956, L. Hubricht (RLH); and Wilson Cave, 3, 9 August 1964, T. C. Barr and S. B. Peck (FSCA). Harrison Co., King’s Cave, 3.5 mi. E Corydon, juv. 6, @, L. Hubricht (RLH). Crawford Co., hillside near Wyandotte Cave, ¢, 2, 26 October 1958, T. C. Barr (FSCA). Monroe Co., Bloomington, 2 ¢, 11 2°, date unknown, C. H. Bollman (USNM) TYPE LOCALITY. Posey Co., upland woods 2.4 mi. N Solitude, 2 ¢, 2 2, L. Hubricht (RLH). Illinois: Union Co., 2.5 mi. NE Alto Pass, Union Point Cave, ¢, 18 August 1968, S. B. Peck (WAS). Monroe Co., Saltpeter Cave, 2 2, 25 June 1965, S. B. Peck (FSCA); and Monroe Cave, Fults, 2, 23 April 1966, S. B. Peck (FSCA). St. Clair Co., Falling Spring, 2, 13 June 1966, S. B. Peck (FSCA). Franklin Co., Zeiglar, 3 6, 2, 20 April 1930, R. V. Chamberlin (FSCA). Missouri: Perry Co., 3 mi. SE Perryville, Lost Cave and Found Cave, 2 6, 2 2, 29 December 1970, D. A. Easterla (WAS); Lost Cave 6, 2, 3 November 1963, R. Stattel (FSCA). Taney Co., Tumbling Creek Cave near Protem, ¢, 10 January 1978, M. W. Fletcher (FSCA). Arkansas: Benton Co., Cave Springs, 6, 2°, 6 April 1951, B. L. Tatum (AMNH). Washington Co., 15 mi. SW Prairie Grove, 6, 3 2, 15 March 1958, M. Hite (RLH); Cave Creek Valley, 6, 4 2, Jan. 1956, M. Hite (FSCA); Devil’s Den State Park, Devil’s Kitchen Cave, juv. 2° , 9 September 1959, J. Tecklin (FSCA); Fayetteville, Mt. Kessler, 9 ¢, 18 2, 4 May 1955, N. B. Causey (FSCA); Ouachita Co., 5 mi. SW Camden, 2 6,2 2, 1 January 1956, A. B. Jones (RLH). Randolph Co., 1 mi. N Pocahontas, ¢, 10 April 1936, L. Hubricht (RVC). Clay Co., Rector, 2 ¢6, 2, Dec. 1951, N. B. Causey (FSCA). Howard Co., sev. 6, sev. 2, 9 December 1953, L. Grey (FSCA). Union Co., Junction City, 6, 3 2, 20 December 1952, N. B. Cau- 562 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON sey (FSCA). Columbia Co., Magnolia, 5 6, many ° 2, 24 December 1949, N. B. Causey (FSCA). Polk Co., 5 N Scottsville, Pine-cedar-hardwood for- est, 2, 6 February 1955, B. Owen (FSCA). Oklahoma: Sequoyah Co., Cottonwood Cave, 6, 25 June 1970, J. H. Black (FSCA). Adair Co., Three Forks Cave between Stillwell and Bunch, juv. 3d, 26 January 1974, J. H. Black (FSCA). Tennessee: Jefferson Co., Indian Cave near Jefferson City, 3 ¢d,2 2, 24 December 1924, J. D. Ives (FSCA); 5 6,6 2, 12 February 1925, J. D. Ives (MCZ); and 3 36, 2 2, 18 April 1925, J. D. Ives (MCZ). Roane Co., Berry Cave,5 46,4 2, 10 July 1963, R. Brandon, J. Huheey (FSCA). DeKalb Co., Avant (Lindsay Williams) Cave, 6, 23 December 1956, T. C. Barr (FSCA). Hamilton Co., Lookout, 2 6, 3 2°, date unknown, Nathan Banks (MCZ). Alabama: Madison Co., 4 mi. E Madison, Matthews Cave, 6 6, 18 2, 2 juvs., 22 August 1968, S. B. Peck (WAS). Colbert Co., McKinney Cave #2, 3 mi. W Sheffield, 2 ¢6, 5 2, 19 December 1965, S. B. Peck (FSCA); and Murvel’s Cave, 4 mi. NW Cherokee, ¢, 2 2, 19 December 1965, S. B. Peck (FSCA). Mississippi: Jeff Davis Co., 3 mi. SSW Prentiss, near White Sand Cr., 2 6, 2 2, 25 December 1959, L. Hubricht (RLH). Franklin Co., 7 mi. W Meadville, 2 6, 5 2, 24 December 1954, N. B. Causey (FSCA). Louisiana: Lincoln Par., Ruston, Pugh’s Pond area, 2, 10 March 1958, W. Harmon (FSCA). Tangipahoa Par., under rotting pine logs in open pine woods, 3 mi. SW Holton, 3 6,4 2, 7 February 1966, K. A. Arnold (FSCA). Natchitoches Par., Creston, 2 6, 18 2, 22 February—5 March 1915, K. Schmidt (MCZ). Cambala hubrichti Hoffman Figs. 6, 11 Cambala annulata (nec Say, 1821) Loomis, 1938:37-38, fig. 11. Chamberlin and Hoffman, 1958:174. Cambala hubrichti Hoffman, 1958:93—94. Type locality.—Bluff along Doe River, 1 mile NW Hampton, Carter Co., Tennessee (Hoffman 1958). Range (Fig. 11).—Southern Appalachian Mountains of southwestern Vir- ginia, eastern Tennessee, and western North Carolina, from Wythe Co., Va., to the Nantahala Gorge, Swain Co., NC (Hoffman 1958). This can now be expanded to include the Black Mountains of NC (NCSM collection) and the Highlands Plateau of Macon Co., NC (one 2 in Hoffman collection). Remarks.—The illustration of the posterior gonopod by Loomis (1938, fig. 11b) is not entirely accurate. I have examined dozens of males and found that the anterior lobe of the coxa is not smoothly rounded but slightly acute VOLUME 92, NUMBER 3 563 anteriodistad, and the posterior lobe is distinctly pointed on the anterior side rather than flattened. Cambala ochra Chamberlin Figs. 8, 12, 13 Cambala ochra Chamberlin, 1942:3, fig. 1. Chamberlin and Hoffman, 1958:175. Cambala saltillona Chamberlin, 1943:3, figs. 1-2; 1952a:30. Chamberlin and Hoffman, 1958:175. NEW SYNONYMY. Troglocambala loomisi Hoffman, 1956:10-11, figs. 9-11. NEW SYNON- YMY. Cambala loomisi Causey, 1964:237. Shear, 1969:135. Type locality.—Darlington, St. Helena Par., Louisiana. Range (Figs. 12, 13).—The distribution of ochra, based solely on male specimens, is from southern Indiana through central Kentucky, and the Cumberland Plateau region of Tennessee to the southern Gulf Coastal Plain, ranging westward into eastern Texas. It is sympatric in most of this area with minor, a fact which renders examination of the posterior gonopods of male cambaloids from their common range mandatory for identifications. Records of the latter species from Louisiana and the Cumberland Plateau by Causey (1959a, 1963) are unreliable and could well be ochra instead. In their common area of occurrence, ochra seems to be more abundant in the southern part and minor, in the northern part. The latter has a much broader range, however, being recorded from Arkansas, West Virginia, and Virginia, where ochra has not been encountered. Remarks.—The type specimen of ochra is immature, and the anterior coxal lobe of the posterior gonopod is incompletely developed. However, the long basal stalk, typical of saltillona and loomisi, is present, and I therefore believe the three names to be synonymous, with ochra having priority. Collections of mature males from St. Helena Par., LA, can confirm this. The configuration of the anterior coxal lobe of the posterior gonopod of ochra is more variable than in any other species of Cambala; the illustration (Fig. 8) is of the most common variant. In some males there is no distal hook, whereas in others it is more pronounced. Some specimens have a larger caudal projection than shown here, which is occasionally subequal to the anterior projection (in males having a reduced distal hook). No geo- graphic pattern is evident to this variation, and, hence, there is no reason to recognize subspecies. The long basal stalk is common to all these variants and thus is the diagnostic feature of the species. 564 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Cambala speobia (Chamberlin) Figs. 9, 13 Eclytus (nec Holmgren, 1855) speobius Chamberlin, 1952a:11. Eclomus speobius Chamberlin, 1952b:71. Chamberlin and Hoffman, 1958:175. Cambala speobia Causey, 1964:243-—244; 1971:273, figs. la, 2a. Cambala caeca Loomis, 1953:417, figs. 1-3. Chamberlin and Hoffman, 1958:174. Cambala captiosa Causey, 1959b:69-71, figs. 1-3. Cambala reddelli reddelli Causey, 1964:239-241, figs. 1-4. NEW SYN- ONYMY. Cambala reddelli inornatus Causey, 1964:241—242, fig. 5. NEW SYNON- YMY. Type locality.—Wyatt Cave, Sonora, Sutton Co., Texas (Chamberlin, 1952a). However Causey (1964) stated that Wyatt Cave was in adjacent Edwards Co. and not Sutton Co. Range (Fig. 13).—Specimens have been examined from Williamson, Tra- vis, Edwards, Culberson, and Wheeler cos., Texas, and the Edwards Pla- teau of Texas in general. Causey (1964) recorded it from 20 miles N Chil- dress, Childress Co., Texas; and west of Cowles, San Miguel Co., and about 15 miles SW Cimarron, Colfax Co., New Mexico. Causey (1971) and Shear (1973) reported it from a cave in Coahuila State, Mexico. Two male cam- baloids that resemble speobia in the configuration of the posterior gonopods have also been collected from Drain, Douglas Co., Oregon (AMNH). Remarks.—Causey (1964) transferred speobius into Cambala and syn- onymized caeca and captiosa. At the same time, however, she proposed two additional forms, reddelli reddelli and reddelli inornatus, which are virturally identical with the holotype of speobia and thus also synonyms. Moreover, I can find no significant differences between the two subspecies of reddelli and see no reason to distinguish the north Texas populations taxonomically. Cambala texana Loomis Figs. 10, 13 Cambala texana Loomis, 1938:40-41, fig. 13. Chamberlin and Hoffman, 1958:175. = Fig. 11. Distribution of annulata (diagonal lines) and hubrichti (cross hatched area). The shaded areas encompass all known, authentic records, including the range extremes in all directions. VOLUME 92, NUMBER 3 ase, sen Oupecibcnce Sia ae RC” saga aoe ea ie ty a L. \ «| “3,8 of : ie \\N¥ (Vag, Ue ge 2 Ett ae PAT te oaien . pesos Gol ALAN ihe VERSES pueneetamen NON, als NN Sy Sa INS WA Ve eS 4 “9 oe 7 REQ SN \ RY SS AEE ORAS IQS EO EA\N\\\ NWA SS Be BP IAW AQUA aS ae C~ PONS \ WW \ BOM EEE WW DUI a»_r : SRY IN SSQn 11 IW QVGA QA AA NY TT ELI AMMA QW ALAS NY Bt BKWAQC$}OWO SNS MA SWS VOWS WY WOWy N \y N ¥ N BN -NMAS S » ~ 8 ‘\ “A Fane ed \A\W\W\ Wy a AWWA AAS A TESTS SQN ATV LI ee NAS eh NES CG LADS a 3 565 566 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Type locality.—Nacogdoches, Nacogdoches Co., Texas. Range.—Known only from the type locality. Remarks.—The illustrations by Loomis (1938), while accurate represen- tations of the characters of the species, are not from the holotype, which was undissected when I received it. On the holotype, the uncate projection of the anterior coxal lobe of the posterior gonopod appears more strongly curved than in Loomis’ illustration. Cambala washingtonensis Causey Cambala washingtonensis Causey, 1954:85, fig. 9. Chamberlin and Hoff- man, 1958:175. Type locality.—Wilma, Garfield Co., Washington. Range.—Known only from type locality. Remarks.—Causey (1954) described this as the ‘‘first western species of the genus’’ from a female, but Chamberlin and Hoffman (1958) noted that it was probably not congeneric with the eastern species of Cambala. In the description of reddelli reddelli, Causey (1964) stated that there is “‘very little difference between the gonopods of C. reddelli and C. washingtonensis.”’ Since to my knowledge no additional material of washingtonensis from the type locality has ever been collected, and the gonopods have certainly never been illustrated or described, it is a mystery how she knew their structure, much less that they were similar to those of reddelli. The wide geographic disparity between the two species would seem to render gonopodal similar- ity unlikely, especially when one notes the considerable differences between the gonopods of ochra, speobia, and texana, all of which occur relatively close together in Texas. In fact, the gonopods of reddelli are virtually iden- tical to those of speobia, and if Causey’s statement is accurate, washing- tonensis must also fall as a synonym of speobia. One wonders if any gon- opods of washingtonensis had been seen at the time this statement was made, and if so, why they were not illustrated to resolve the question of the generic position of this species. Additionally, Causey (1964) reported the range of Cambala as including ‘‘northern Idaho,’’ but no specific Idaho records have ever been reported, and no such specimens are available in any museum collection. Consequently, one must assume that the basis for this citation of Idaho was the location of washingtonensis in an adjacent part of Washington. Some insight into the washingtonensis problem is now possible due to the two male specimens of Cambala from Drain, Douglas Co., Oregon, which I discovered among unsorted material at the AMNH. As indicated earlier, the posterior gonopods are similar to those of speobia, suggesting that washingtonensis may indeed be a species of Cambala and a synonym of speobia. Unfortunately, Douglas Co. is in southwestern Oregon and sev- VOLUME 92, NUMBER 3 567 rH ae Sr q Peas A aa ) ean Aawa Orage =velbage, — Se rt ey aN Gce= ne) — BY A ay SP Geen “eel Pet BNO OG As: tt UH fee a ieeaeateees Sees | ee Texfea feat | Be GY? ake Sos Gagu CoN >) A eS TAL wae 1S tess, aA A AY SeBET eta ceri enero ine cae ee EE a ae oe ARO OTL STIS A CPages CH A Bie BS bers lars ce SNS eee), ALESS Per PONT OT RR WO RAR TED ERS I LEPC Ln Pree ES EE) Pade eR S SGN CEE ER Att Spare TES iz oT AO tae eT Bape toh areuteee i careaiec Snes celta on oe Py Ee ee Sy EL EL Tila Tad OP EORTC Ye yo 5 < SMI eer ree te ier TS Fekete Ame SOO Seeautaw reese tea eas BLS Vn) ae L Lae ia “EEE ain] Norwtaab Sees) Fr VES yd nepecee Bea ati ealee Minn, TT SSS BOT CREP Ee rec Pareto Stal ctnie ait cence TETAS XIE i far ALLIES POR US ar Shs ea Sep eT RRs A NB AOR OY ¥ TX ‘ans re oq CIS ITN Hive: m1 AN ez Gwe oak ae =< ny; Fig. 12. Distribution of minor (circles) and ochra (triangles). eral hundred miles from Garfield Co., Washington, which is in the south- eastern corner of that state. Hence, it would be risky to assume that the Oregon material is conspecific with that from the latter area, and the Oregon locality is reported merely for the sake of completion. Specific identification of this material should await collection of male topotypes of washingtonen- sis, but they do establish the presence of Cambala in the Pacific northwest. Causey (1964) may have been correct in stating that the gonopods of wash- ingtonensis closely resemble those of reddelli (which would make washing- tonensis a Synonym of speobia), but the basis for this statement is obscure since the type and only reported specimen is a female. As noted in the introduction, the type of washingtonensis is absent from its published re- pository, the AMNH, so I was unable to examine it personally, and no 568 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON material identified as washingtonensis was found in the Causey collection (now at FSCA). A Note on the Occurrence of Cambala in Caves Perhaps the most intriguing aspect of the biology of Cambala is the oc- currence of many of its species in caves. To date, hubrichti, texana, and washingtonensis, the last two known only from their type localities, have never been reported from a subterranean habitat, and I can now remove hubrichti from this list. During a recent survey of North Carolina caves by personnel of the NCSM, males and females of hubrichti were collected from caves in McDowell, Polk, Rutherford, and Transylvania cos. Specimens were found in the dark zone of Bat Cave, Rutherford Co., and the aphotic zone of Limekin Cave, McDowell Co. To the best of my knowledge, this is also the first report of cave millipeds from North Carolina. The other species of Cambala commonly inhabit caves. The type species, which is distinct enough to render reliability to literature records, has been reported from caves in Alabama and Florida (Loomis 1943) and Tennessee (Causey 1959a). To these can now be added Wyandotte Cave, Crawford Co., Indiana (one male in MCZ collection) and Carter Cave, Robertson Co., Kentucky (one male in FSCA collection). Causey (1959a) reported minor as well as annulata from several Tennessee caves, but some of these records may be for ochra instead. Most specimens of ‘‘minor’’ were found on bat, cave rat, or raccoon dung in moist areas in the caves. She recognized three groups of cave millipeds in Tennessee—accidental species, including an- nulata, ““epigean species that are somewhat tolerant of cave conditions”’ and hence occur there more often than those in the first category, and troglobites. Cambala minor was included in the second group. In a paper on cave millipeds of Missouri, Causey (1960) noted the wide geographic distribution of minor and suggested that it probably occurred in many caves of that state. This prediction is confirmed herein, as male specimens of minor have been examined from caves in Perry and Taney cos., Missouri. Due to the confused nomenclature of minor, only a few cave records of ochra exist which can be considered reliable. One, involving the type ma- terial of Troglocambala loomisi, a synonym of ochra, is from Turk’s (Brooklyn) Cave, Conecuh Co., Alabama. Shear (1969) noted the identity problem with many cave records of minor and classified both minor and annulata as being troglophilic, e.g., animals that occur in caves and can complete their life cycle there, but lack modi- fications for this life and also live in epigean habitats. This conforms to the second of Causey’s (1959a) categories, and the two authors differ in the position of annulata. Shear also suggested that speobia in Texas may be a true troglobite, e.g., an animal occurring solely underground, usually with VOLUME 92, NUMBER 3 569 Fig. 13. Distribution of speobia (circles), texana (triangle), and ochra (squares) in Texas; the open symbol indicates a literature record believed to be valid. On the inset map, a smooth curve has been drawn around all records of Cambala to show the generic range. Oklahoma and southern Kansas are included even though no material has been collected from these states. The type locality of washingtonensis and that of the specimens from Oregon are shown in the northwest. modifications for this life, and indeed this seems to be the case. All records of speobia or its synonyms known to me are from caves in Texas, New Mexico, and Mexico; the milliped has never been encountered in a surface habitat. Causey (1964, 1971) considered it a troglobite and noted the mod- ified legs and antennae, and the absence of ocelli. Thus, although annulata, minor, ochra, and speobia all occur in caves, they are significantly different in the frequency of their occurrence there and the degree of their modifi- cation for subterranean life. Literature Cited Bollman, Charles H. 1887. Notes on North American Julidae.—Ann. N. Y. Acad. Sci., 4:25—44. . 1888a. Catalogue of the myriapods of Indiana.—Proc. U.S. Natl. Mus., 11:403-410. 570 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON . 1888b. Notes on a collection of Myriapoda from Mossy Creek, Tenn., with a descrip- tion of a new species.—Proc. U. S. Natl. Mus., 11:339-342. 1888c. A preliminary list of the Myriapoda of Arkansas with descriptions of new species.—Ent. Amer., 4:1-8. . 1893. The Myriapoda of North America.—U. S. Natl. Mus. Bull. No. 46, 210 pp. Brimley, C. S. 1938. Insects of North Carolina.—North Carolina Department of Agriculture, Division of Entomology, Raleigh, 560 pp. Causey, Nell B. 1953. On five new North American millipeds and records of some established species.—Am. Mid. Nat., 50:152—158. . 1954. The millipeds collected in the Pacific northwest by Dr. M. H. Hatch.—Ann. Ent. Soc. Am., 47:81-86. . 1959a. Some cavernicolous millipeds from the Cumberland Plateau.—J. Tenn. Acad. Sci., 34:229-237. ——. 1959b. Two new troglobytic millipeds from Texas.—Proc. Biol. Soc. Wash., 72:69-74. . 1960. Troglobitic millipeds in Missouri.—Missouri Speleol., 5:60-65. . 1963. Additional records of Louisiana millipeds.—Proc. La. Acad. Sci., 26:76-79. . 1964. New cavernicolous millipeds of the family Cambalidae from Texas and New Mexico.—Int. J. Speleol., 1:237—248. . 1971. The Cambalidae in Mexican caves, with descriptions of three new species of Mexicambala (Diplopoda: Cambalidae).—Proc. Biol. Soc. Wash., 84:271—282. Chamberlin, Ralph V. 1918. Myriapods from Nashville, Tennessee.—Psyche, 25:23-30. . 1942. New southern millipeds.—Bull. Univ. Utah Biol. Ser., 6(8):1-19. . 1943. On some genera and species of American millipeds.—Bull. Univ. Utah Biol. Ser., 8(2): 1-20. 1947. Some records and descriptions of diplopods chiefly in the collection of the Academy.—Proc. Acad. Nat. Sci. Phila., 99:21—58. . 1952a. Three cave-dwelling millipeds.—Ent. News, 63:10-12. . 1952b. Eclomus, nom. nov. (Diplopoda).—Ent. News, 63:71. . 1952c. Further records and descriptions of American millipeds.—Great Basin Nat., 12:13-34. , and Richard L. Hoffman. 1958. Checklist of the millipeds of North America.—U. S. Natl. Mus. Bull. No. 212, 236 pp. Cope, Edward D. 1869. Synopsis of the extinct Mammalia of the cave formations in the United States, with observations on some Myriapoda found in and near the same, and on some extinct mammals of the caves of Anguilla, W. I., and other localities.—Proc. Am. Philos. Soc., 11:171-192. Gervais, P. 1837. Etudes pour servir a l’histoire naturelle des Myriapodes.—Ann. Sci. Nat., Zool., Ser. 2, 7:35-60. Gray, John E. 1832. Myriapods, Jn: Griffith, The Animal Kingdom Arranged in Conformity with its Organization by the Baron Cuvier, 15 (Class Insecta, vol. 2):784, pl. 135, figs. 2, 2a—c; Whittaker, Treacher & Co., London. Hoffman, Richard L. 1956. New genera and species of cavernicolous diplopods from Ala- bama.—Geol. Surv. Ala., Mus. Pap. No. 35, 13 pp. . 1958. Appalachian Cambalidae: Taxonomy and distribution (Diplopoda: Spirostrep- tida).—J. Wash. Acad. Sci., 48:90—94. Loomis, Harold F. 1938. The cambaloid millipeds of the United States, including a family new to the fauna and new genera and species.—Proc. U.S. Natl. Mus., 86:27—66. . 1939. The millipeds collected in Appalachian caves by Mr. Kenneth Dearolf.—Bull. Mus. Comp. Zool., 86:165—193. VOLUME 92, NUMBER 3 Swi 1943. New cave and epigean millipeds of the United States, with notes on some established species.—Bull. Mus. Comp. Zool., 92:371-410. . 1953. New millipeds of the western states and lower California.—J. Wash. Acad. Sci., 43:417-422. McNeill, Jerome. 1888. A list, with brief descriptions of all the species, including one new to science, of Myriapoda of Franklin Co., Ind.—Bull. Brookville Soc. Nat. Hist., 3: 1-20. Say, Thomas. 1821. Descriptions of the Myriapodae of the United States.—J. Acad. Nat. Sci. Phila., 2:102—114. Shear, William A. 1969. A synopsis of the cave millipeds of the United States, with an illustrated key to genera.—Psyche, 76:126—-143. 1973. Millipeds (Diplopoda) from Mexican and Guatamalan caves.—Proc. Accad. Naz. dei Lincei, 171:239-305. Shelley, Rowland M. 1977. Appendicular abnormalities in the milliped family Xystodesmidae (Polydesmidae).—Can. J. Zool., 55:1014—-1018. . 1978. Millipeds of the eastern Piedmont region of North Carolina, U.S.A. (Diplopo- da)i——S. Nat. Hist... 12537-79: Williams, Stephen R., and Robert A. Hefner. 1928. The millipedes and centipedes of Ohio.— Bull. Ohio Biol. Surv., 4:93-146. Wood, Horatio C. 1865. The Myriapoda of North America.—Trans. Am. Philos. Soc., 13:137- 248. Wray, David L. 1967. Insects of North Carolina, Third Supplement.—North Carolina De- partment of Agriculture, Division of Entomology, Raleigh, 181 pp. North Carolina State Museum, P.O. Box 27647, Raleigh, North Carolina 27611. PROC. BIOL. SOC. WASH. 92(3), 1979, pp. 572-576 NOTE ON THE OCCURRENCE OF A SWARM OF SALPA CYLINDRICA CUVIER (TUNICATA: SALPIDAE) IN SARDINERA LAGOON, PUERTO RICO Takasi Tokioka and P. B. Bhavanarayana Abstract.—The occurrence of a dense swarm of solitary individuals of Salpa cylindrica Cuvier in Puerto Rican waters is reported, and evidence is presented that solitary individuals of Jasis zonaria (Pallas) may produce similar swarms in Japanese waters. Morphological details of the Puerto Ri- can specimens are described and illustrations provided. A sample from a swarm of a pelagic tunicate observed by Mr. T. A. Wiewandt during the first week of July 1974 in Sardinera Lagoon, Isla de Mona, Puerto Rico, was forwarded to the authors for identification by Dr. F. M. Bayer of the National Museum of Natural History, Smithsonian In- stitution. The specimens were solitary individuals of the tropical salp, Salpa cylindrica Cuvier, 1804. According to notes of the observer, these salps occurred in a swarm of thousands (millions?) of individuals that looked in life like clear pulsating cylinders each with a solid blue sphere at one end; further, a similar swarm had been seen, also in brief abundance, at Playa de Pajaros two years before (1972). The notes seem to imply that the entire swarm, like the sample examined, was composed solely of solitary individ- uals of S. cylindrica. This is very interesting, because swarms of salps are generally composed of a larger number of aggregate forms and a much smaller number of solitary forms. This proportion is quite natural, since asexual propagation by budding of the stolon in the solitary forms is incom- parably greater than the sexual propagation by placental embryos in the aggregate forms. Thus, the formation of dense swarms by only the solitary forms must be rather rare, though not impossible. However, one of the au- thors has found that the stomachs of a number of mackerels from Japanese waters were full solely of the solitary forms of Jasis zonaria (Pallas). As the aggregate form of that salp is nearly as large as the solitary form, no feeding selection is possible by the fish. This observation is evidence for the for- mation of swarms in Jasis consisting only of solitary forms. Occurrences of swarms of only solitary forms in salps indicate both well synchronized breeding and some stability of the water masses inhabited by such swarms, although they might be formed by accumulation on the bound- aries between water masses. If the latter is the case, it follows that some evidence of morphological change leading toward speciation among popu- VOLUME 92, NUMBER 3 573 Fig. 1. Stolon with narrow distal part (fd). a, 18 mm long specimen; b, 23 mm long speci- men, c, 16 mm long specimen. lations of those salps separated from one another in different water masses may be expected. In anticipation of this, some detailed observations on the specimens subjected to examination, especially on the numbers of muscular bands in the respective body muscles as recorded in Table 1, were made to help future studies. In all of the 14 specimens examined, from 16 mm to about 24 mm in length, the nucleus is situated between body muscle VIII and cloacal muscle PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 574 ‘poyeredas ureulol jnq si9yj0 oy} Yoeoidde ‘ystio\se ue Aq UMOYS ‘Sa[dSNUI 9S9y} JO SWOS ‘I9AQMOY ‘SUSUTIDEdS OWOS UT “1OYyJOUe 9UO WIM AT[esIOppIu }DejU0D UI AT[eIOUNs oe AJ-] Sojosnw Apog , se /EE| SIZ L6l SLI 861 €Ll 8LI ps €8I L9I 6LI ILI v6 y9l [R101 8°91 is 61 81 81 6 91 91 LI LI vl bl LI 61 SI J 9f9snu [eseo[) 9°07 VC Sé v7 8 GC LI v7 (K6 02 cl (G6 81 6 SI IIIA Ld oC 87 a6 €C 76 8I £7 61 ST 61 07 91 6 IZ IIA 02 I? €¢ IZ 81 v7 07 61 CC I? 61 07 bl (G6 91 IA 8°77 Sc 97 v7 (G6 97 (G6 61 vC EG 02 (G6 CC €C IZ A LI ¥VT +97 v7 +81 IZ +07 €C 61 07 +07 6 #17 #€C (G6 AI 9°07 +61 *€C IZ IZ LI v7 LI 02 €C *CC 07 + V7 WG LI Ill CsI 61 eC IZ 81 6 81 91 81 91 02 81 *S 1 81 91 II IZ (G6 (G6 IZ 61 EG 81 17 €~ 81 81 02 v7 vc IZ J 9J9snw Apog ues, +WW 07 WW pz WUE¢Z WW7Z WW]Z WW{Z WWOQZ Wwe? WUQZ WU 6] WWE] WU Eg] WwW] WU 9g] yysue, Apog vl el Gli IT Ol 6 8 IE 9 ¢ 14 € c I ‘ON uouIDedg ‘soposnul Apoq Ul spueq Je[NosnuI Jo UONNGMSIg “| IIqeL VOLUME 92, NUMBER 3 x75 2 cl.m.f N V4 clmJI —————————————————— SS ae Fig. 2. Sinus (si) connecting ends of posterior branch of cloacal muscle I (cl.m. I) to mid- ventral part of cloacal muscle II (cl.m. II), 23 mm long specimen. I (defined by some specialists as body muscle IX). In all the specimens, the stolon generally reaches anteriorly to body muscle VI; in some specimens the distal part of the stolon is narrower than the subdistal part because the stolon-individuals are less developed in the former than in the latter (Fig. 1), and this narrow distal part may extend anteriorly to body muscle V or turn back to the middle of the space between body muscles VI and VII. It is not known, however, whether or not such a distal structure of the stolon is usual, at least in young (or small) solitary forms of the species from the sea area concerned. Cloacal muscle I is divided ventrally into anterior and posterior branches. The anterior branch is mid-ventrally divided clearly, with each end terminating near the posterior end of the stomach, and con- sists of 8 to 13 muscular bands; while the posterior branch consists of 5 to 9 muscular bands and is divided mid-ventrally rather indistinctly at a level slightly posterior to the anus, usually with a sinus connecting both ends of the branch to the ventral middle of cloacal muscle II; this shows that the posterior branch might originally be in contact with cloacal muscle II in the mid-ventral line (Fig. 2). Cloacal muscle II consists of 9 to 14 muscular bands, the first atrial sphincter of about 6, the second of about 3, and the third of 2 muscular bands followed by up to 8 fine sphincters each consisting of a single band. Acknowledgment Before closing the present short note the authors wish to express their cordial thanks to Dr. Bayer for the opportunity of examining these speci- mens from the West Indies. Literature Cited *Fagetti Guaita, Elda. 1959. Salpas colectadas frente a las costas central y norte de Chile.— Rev. Biol. Mar. Valparaiso 9(1—3):201-228, pls. 1-6. * In these papers, Salpa cylindrica is referred to as Weelia cylindrica. 576 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Metcalf, M. M. 1918. The Salpidae: a taxonomic study.—U.S. Nat. Mus. Bull. 100, vol. 2, part 3:3-193, pls. 1-14. Sewell, R. B.S. 1953. The pelagic Tunicata.—John Murray Exped. 1933-34. Sci. Rep. 10(1):1- 90, 1 pl., 32 figs. Thompson, Harold. 1948. Pelagic tunicates of Australia——Commonwealth Council for Sci- entific and Industrial Research, Australia, Melbourne. 196 pp., 75 pls. *Yount, J. L. 1954. The taxonomy of the Salpidae (Tunicata) of the Central Pacific Ocean.— Pacific Sci. 8:276—330. (TT) Rinkai 454-3, Shirahama, Wakayama-ken, Japan 649-22; (PBB) An- dhra University, Waltair, India. PROC. BIOL. SOC. WASH. 92(3), 1979, pp. 577-579 LOUISIANA AND PANAMA CANAL LOCATIONS AND ECOLOGY OF MUNNA (PANGAMUNNA NOV. SUBGEN.) REYNOLDSI FRANKENBERG & MENZIES (ISOPODA: ASELLOTA) George A. Schultz Abstract.—Specimens of what were formerly called Munna (Uromunna) reynoldsi Frankenberg & Menzies (1966) (family Munnidae), were collected in the brackish water of Lake Pontchartrain, New Orleans, Louisiana, and in the fresh water of the locks of the Panama Canal. The species is placed in a new subgenus, Pangamunna, based on the fact that no mandibular palps are present. The specimens from the canal are the first munnids from apparently established populations in fresh water. Number in each sex, lengths and embryo number in the marsupia of some females are included in the discussion. Specimens of tiny isopod crustaceans from Louisiana and Panama were examined by the author and identified as Munna (Uromunna) reynoldsi Frankenberg & Menzies (1966) (family Munnidae). The species was de- scribed originally from oak leaves in salt marshes on Sapelo Island, Georgia. The type-specimens were collected in tidal regions where brackish and per- haps fresh water are found from time to time. The specimens discussed here are from Lake Pontchartrain, Louisiana and the Panama Canal. The canal locations are from the locks on the Atlantic (Gatun) and the Pacific (Mir- aflores and Pedro Miguel) slopes of the canal. The species as described by Frankenberg & Menzies lacks mandibular palps. The specimens from the two locations also lack palps (Fig. 1). All species placed in the subgenus Uromunna Menzies (1962:36) and M. (U.) magnifica Schultz (1964) have triarticulate mandibular palps. The author thinks that lack of palps is a character which is of at least as much value for defining a subgenus of Munna Kr@yer as is the shape and spination of the rami of the uropod. A new subgenus, Pangamunna (Panama-Georgia-Munna combined— gender feminine) with M. (U.) reynoldsi as type-species, is thus proposed here. It has the characters of Munna Kr@yer, but lacks mandibular palps. So far no other species are known since the presence or absence of man- dibular palps has not been recorded for some species of Munna. When more species are discovered the definition of the subgenus might be expanded to 578 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Figs. 1-3. Munna (Pangamunna) reynoldsi: 1, Right mandible; 2, Apex of fused male pleopods 1; 3, Uropod. include characters other than the lack of palps. The genus Munnogonium George & Stromberg of the Munnidae also lacks mandibular palps (Bowman & Schultz, 1974), but it is clearly not related to Pangamunna. M. reynoldsi is also the only genus of Munna to be taken from fresh water in the New World. The specimens from the locks of the Panama Canal examined here (both males and females) correspond in morphology remarkable well with the description of M. (P.) reynoldsi. Eight spines are present on the apex of the fused male pleopods 1 (Fig. 2). Antennae 1 and 2 are similar, and pe- raeopods I and VII (male) are also similar. The uropod is illustrated here (Fig. 3). The smaller ramus was not seen even though the posterior part of several specimens were dissected and examined. Other minor differences include number of coupling hooks on the endite of the maxilliped and the pattern of spination on various parts of the body. The specimens from the canal have from 2 to 4 coupling hooks; M. (P.) reynoldsi from Georgia has 3. The pattern of spines on the maxillipedal palp, on the posterior margin of peraeopod I and on the apex of male pleopod 2 differ slightly. The 47 specimens from the locks range in length from 0.7 to 1.3 mm. Males (17) are slightly shorter than females (30). Nine females were gravid. The number of embryos in undisturbed marsupia ranged from 6 to 10. Males and gravid females were present at each lock. In June 1973 (4 females) and during July and August 1975 specimens (133 males and females) were taken at Lake Pontchartrain, New Orleans, Louisiana. The greatest number of specimens was taken at the mouth of the inner harbor navigation canal. Mandibular palps were absent. Setation on the apex of male pleopods | and other pleopods, flagellar article number on antennae and general characters VOLUME 92, NUMBER 3 579 of the peraeopods and uropods were examined and also confirmed that the specimens were M. (P.) reynoldsi. Gravid females were present. Salinity of the water at the principle collection location varied from 2 to 12%. during the time of collection. The existence in the fresh water of an isopod whose relatives are from marine and brackish waters has interesting zoogeographical implications. The spread of marine animals through the Panama Canal has for the most part been limited by the freshwater barrier created by Gatun Lake. How- ever, M. (P.) reynoldsi is able to live and apparently breed in fresh water so the lake is not a barrier to its distribution. Unfortunately, the species has not been collected in the brackish or salt water at either end of the canal so whether or not it has extended its range through the canal to the Pacific Ocean is not known. All species of Munna recorded so far on the Pacific coast (in Chile—Menzies, 1962; in California—Schultz, 1964) have triartic- ulate mandibular palps. Acknowledgments The author would. like to thank Dr. Meredith L. Jones, Division of Worms, National Museum of Natural History, for the specimens from the Panama Canal, and Dr. Michael Poirrier, Department of Biology, University of New Orleans, for the specimens from Louisiana. Specimens from the canal are deposited in the collections of the National Museum of Natural History. Literature Cited Bowman, T. E., and G. A. Schultz. 1974. The isopod crustacean genus Munnogonium George and Stromberg, 1968 (Munnidae, Asellota).—Proc. Biol. Soc. Washington 87(25):265-— 27. Frankenberg, D., and R. J. Menzies. 1966. A new species of asellote marine isopod, Munna (Uromunna) reynoldsi (Crustacea: Isopoda).—Bull. Mar. Sci. 16(2):200—208. Menzies, R. J. 1962. The zoogeography, ecology and systematics of the Chilean marine iso- pods.—Lunds Univ. Arsskr., N.F. Avd. 2, 57(11):1-162. Schultz, G. A. 1964. Some marine isopod crustaceans from off the southern California coast.— Sci. 18(3):307-314. 15 Smith St., Hampton, N. J. 08827. PROC. BIOL. SOC. WASH. 92(3), 1979, pp. 580-588 STENOCOROPHIUM BOWMANI, A NEW GENUS AND SPECIES OF THE FAMILY COROPHIIDAE FROM THE PALAU ISLANDS (CRUSTACEA: AMPHIPODA) Gordan S. Karaman! Abstract.—A new genus and species of the family Corophiidae, Steno- corophium bowmani is described from Babelthuap Island (Palau Islands in southern Pacific, E. of Philippines). This genus is closely related to the genus Paracorophium Stebbing 1899, but differs in the simple ¢ gnathopod 2 and the uniramous uropod 3. Thanks to Dr. Thomas E. Bowman and Dr. J. Laurens Barnard from the Smithsonian Institution, Washington, D.C., I was able to study an amphipod from the bottom of the Ngermeskang River, above tidal influence, on Ba- belthuap Island (Palau Islands in the South Pacific). The specimen proved to represent a new genus and species of the family Corophiidae, described below. Stenocorophium, new genus Diagnosis.—Body smooth, Paracorophium-like, urosomites 1-2 co- alesced, urosomite 3 free. Antennae normal; accessory flagellum absent. Labrum incised symmetrically; labium with long inner lobes. Maxilla 1: inner lobe short, outer lobe with ca. 7 spines, palp 2-articulate. Maxilla 2: both lobes moderately narrow, inner lobe without dorsal oblique row of setae. Maxilliped: outer lobe with a row of slender spines along inferior margin; palp 4-articulate, palp article 2 elongated, palp article 3 not lobed. Mandible: incisor toothed, molar triturative, palp 3-articulate. Coxa | produced at ventroanterior part, coxa 4 without distoposterior lobe, coxa 5 long. Gnath- opod | subchelate. Gnathopod 2 simple, but its article 4 elongated and article 5 attached proximally on article 4, like that in genus Paracorophium. Article 2 of pereopods 6-7 lobed, pereopod 7 extremely enlarged. Uropods 1-2 well developed; peduncle of uropod 1 with ventrodistal long tooth like that in genus Paracorophium. Uropod 3 short; inner ramus absent; outer ramus uniarticulate, as long as peduncle. Telson fleshy, short, entire, with distal corner tooth on each side in lateral view. Coxal gills ovoid, normal. Type-species.—Stenocorophium bowmani, n. sp. 1 Contribution to the knowledge of the Amphipoda 99. VOLUME 92, NUMBER 3 581 Etymology.—From the Greek, ‘“‘stenos’’ (=narrow, close), and the ge- neric name Corophium. Remarks.—Stenocorophium shows clear affinities with Paracorophium Stebbing 1899 (body-shape, coxae, article 4 of gnathopod 2, uropods 1-2, epimeral plates, pereopods 3-5, gnathopod |, most mouthparts, etc.). Par- acorophium is known from Australia and New Zealand, in brackish waters. Stenocorophium differs from Paracorophium by simple gnathopod 2 in males (subchelate or chelate in Paracorophium), by the presence of only 7 distal spines on outer lobe of maxilla 1 (9 spines in Paracorophium), by the uniramous uropod 3 (biramous in Paracorophium), by the lobed article 2 of pereopods 6—7 (unlobed in Paracorophium). Based on Barnard’s (1973) revision of the family Corophiidae, the rather isolated position of the genera Paracorophium and Stenocorophium in the family Corophiidae is evident, based on the shape of gnathopod 2 in males and females. Stenocorophium bowmani, new species Figs. 1-5 Description.—Male 3 mm (holotype). Body slender, slightly dorsoven- trally compressed, smooth (Fig. 1), urosomites 1—2 coalesced, urosomite 3 free; all 3 metasomites and urosomites 1—2 with 2 dorsal setae each (Fig. 1; 2E-F). Rostrum short, lateral cephalic lobes acute, ventroanterior sinus present (Fig. 4A), eyes ovoid, horizontal. Antennae 1-2 slender, shorter than half body length (Fig. 1). Antenna 1: peduncle articles 1-3 progressively shorter, poorly setose; main flagellum 9-articulate, 3 distal articles with one aesthetasc each; accessory flagellum absent (Fig. 2A). Antenna 2 shorter than antenna 1: peduncle article 3 short, articles 4—5 slender, article 4 longer than article 5, each with 2-3 bunches of setae longer than diameter of articles (Fig. 2A); flagellum 4-articulate, shorter than pe- duncle, bearing several setae longer than diameter of articles, calceolae absent. Antennal gland cone short, not reaching tip of peduncle article 3. Labrum broader than long, symmetrically incised (Fig. 4B). Labium not gaping, with long inner lobes (Fig. 3A); outer lobes without distinct prox- imal dilatations (=fingers). Maxilla 1: inner lobe very small, without setae (Fig. 4C), outer lobe with 7 distal spines having 1-2 lateral teeth each; left and right palp symmetrical to each other, 2-articulate, provided with 5—6 distal slender spines. Maxilla 2: both lobes longer than broad; inner lobe shorter than outer, bearing rows of marginal setae only, dorsal oblique row of setae absent; outer lobe with distal setae only (Fig. 2B). 582 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON i ae [] \ WY My Fig. 1. Stenocorophium bowmani: male, lateral view. Maxilliped: inner lobe with a row of marginal plumose setae; outer lobe bearing a row of slender spines along inferior margin; palp 4-articulate, article 2 elongate, article 3 short, not lobed, article 4 short with distal spines (Fig. SA). Mandible: incisor toothed, molar triturative, strong (Fig. 3B); palp 3-seg- mented: article 1 short, smooth, article 2 with 3 setae; article 3 shorter than article 2, bearing 6 D-setae, 4-5 submarginal C-setae, 2 long E-setae, one group of A-setae and one group of B-setae. Coxae moderate, coxae 1-5 of subequal size, coxae 6—7 short. Coxae 1- 7 with simple or plumose ventral setae (Fig. 2C—D, 4D). Coxa 1 with pro- duced ventroanterior part (Fig. 3C), coxa 4 without any trace of ventropos- _ terior lobe (Fig. 2D). Coxa 5 bilobed, anterior lobe much longer than pos- terior (Fig. 5B); coxa 6 short, bilobed (Fig. 5C); coxa 7 ovoid, entire (Fig. 5D). Gnathopod 1 subchelate (Fig. 3C—D); articles 3—4 short; article 3 with 3 strong and long distoposterior plumose setae; article 5 inflated, bearing nu- merous setae along posterior margin; article 6 shorter than 5, twice longer than broad, with concave posterior margin, poorly setose; palm convex, poorly inclinated, weakly crenellated, without corner or subcorner spines (Fig. 3D); dactyl exceeding width of article 6, with one seta at outer margin. VOLUME 92, NUMBER 3 583 Fig. 2. Stenocorophium bowmani: A, Antennae 1-2; B, Maxilla 2; C, Pereopod 3; D, Pe- reopod 4; E, Urosome and uropods, lateral view; F, Urosome with uropods and telson, dorsal view; G, Distal part of telson, lateral. 584 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 3. Stenocorophium bowmani: A, Labium; B, Mandible; C-D, Gnathopod 1; E, Epi- meral plates 1-3. VOLUME 92, NUMBER 3 585 Gnathopod 2 (Fig. 4D) merochelate, distally simple; article 3 short, article 4 elongate, provided with a row of marginal and submarginal long setae at anterior margin; article 5 elongate, as long as article 4, attached in proximal part of article 4 and forming with it a chela; article 6 reaching nearly half of article 5, narrow, bearing several distal setae only (Fig. 4D); dactyl absent. Pereopods 3-4 alike, with stout articles; article 5 with several longer setae at posterior margin; article 6 longer than 5, tapering distally; dactyl strong, longer than half of article 6 (Fig. 2C—D). Pereopods 5-7 progressively longer. Pereopods 5-6 stout, with articles 3— 6 short, articles 2—5 bearing plumose setae along anterior margin (Fig. 5B- C), dactyl short. Article 2 of pereopod 5 tapering distally, not lobed, with a row of marginal and submarginal plumose setae at posterior margin (Fig. 5B). Article 2 of pereopod 6 oval, not tapering distally, with distoposterior lobe, and with a row of plumose setae along posterior margin (Fig. 5C). Pereopod 7 extremely enlarged (Fig. 5D). Article 2 oval, with distinct dis- toposterior lobe; article 3 short; article 4 large, with distoanterior finger; article S half of article 4, produced anteriorly; article 6 narrow, longer than article 5, bearing short distal dactyl. Pleopods well developed, rami multiarticulate, inner ramus distinctly longer than outer; peduncles with 2 retinaculae each. Epimeral plate 1 remarkably shorter than plates 2-3, all 3 epimeral plates with subrounded ventroposterior margin (Fig. 3E); epimeral plate 1 with one facial plumose seta, plate 2 with numerous facial and ventral plumose setae, plate 3 without facial setae. Posterior margin of epimeral plate 3 convex. Uropods 1-2 well developed (Fig. 2E). Uropod 1: peduncle longer than rami, with dorsal spines, and with long ventrodistal tooth, ventrofacial spine absent; rami with lateral (dorsal) and distal spines, inner ramus longer than outer. Uropod 2: peduncle nearly as long as inner ramus, inner ramus long- er than outer. Uropod 3 short, not reaching tip of uropods 1-2: peduncle short, nearly as long as broad, with one distal spine on the place of inner ramus. Inner ramus absent. Outer ramus nearly as long as peduncle, bearing several distal setae (Fig. 2F). Telson short, fleshy, broader than long, distally with 2 corner teeth re- curved in lateral view (Fig. 2E, F, G), dorsally with 2 simple and 2 short plumose setae. Coxal gills ovoid, simple, on thoracic segments 2-6. Material examined.—Sta. 1 NMK 4-02, silt-covered rocks on bottom of Ngermeskang River, Babelthuap (=Babeldoab) Island, above tidal influ- ence, 27 Jan. 1976, leg. Greg R. Bright, Palau Islands, southern Pacific E. of Philippines, 5 holotype, 3 mm, deposited in the National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560. (USNM 331543) 586 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 4. Stenocorophium bowmani: A, Head; B. Labrum; C, Maxilla 1; D, Gnathopod 2; E, Left uropod 3; F, Right uropod 3. VOLUME 92, NUMBER 3 587 Fig. 5. Stenocorophium bowmani: A, Maxilliped; B, Pereopod 5; C, Pereopod 6; D, Pe- reopod 7. 588 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Etymology.—Named for my colleague, Dr. Thomas E. Bowman. Acknowledgments I thank Dr. Thomas E. Bowman and Dr. J. Laurens Barnard of the Na- tional Museum of Natural History, Smithsonian Institution, Washington, D.C., and the collector, Mr. Greg R. Bright, for making available the ma- terial used in this study. Literature Cited Barnard, J. L. 1973. Revision of Corophiidae and related families (Amphipoda).—Smithsonian Contributions to Zoology 151:1-27. Stebbing, T. R. R. 1899. On the true Podocerus and some new genera of amphipods.—Annals and Magazine of Natural History, London 7(3):237—241. Biological Institute, P.O. Box 40, Titograd, Yugoslavia. PROC. BIOL. SOC. WASH. 92(3), 1979, pp. 589-600 A NEW SPECIES OF PRIMARY BURROWING CRAYFISH OF THE GENUS CAMBARUS FROM THE RIDGE AND VALLEY PROVINCE IN TENNESSEE Raymond W. Bouchard and David A. Etnier Abstract.—Cambarus (Depressicambarus) deweesae, a new species of primary burrowing crayfish, is described from the Ridge and Valley province in Anderson and Roane counties, Tennessee (Tennessee River basin). The known localities lie near the foot of Walden Ridge, an escarpment that delimits the eastern margin of the Cumberland Mountains and Cumberland Plateau sections of the Appalachian Plateaus province. Color notes describ- ing its reddish color pattern, relationships with its closest ally C. (D.) stria- tus Hay, distribution, life history notes and ecological data are presented. The burrowing habits of North American crayfishes were noted as early as 1817 by Constantine S. Rafinesque in his description of Cambarus fossor (suppressed senior synonym of C. (Lacunicambarus) diogenes Girard (1852:88), see Hobbs, 1967:126). The fossorial Cambarus diogenes and chimneys marking its burrows were illustrated in the early 1800’s by John James Audubon in a portrait of the white ibis. He later included a description of the methodology employed by this bird to extract crayfish from burrows. Despite an early knowledge of burrowing crayfishes (the first epigean species from North America was described by Fabricius in 1798), our knowl- edge concerning life history aspects and distributions of many primary bur- rowers remains incomplete. A number of new species of primary burrowers, all with limited ranges, have been described recently (see Hobbs, 1973:463, 469; Hobbs, 1975:24, 28; Schuster, 1976:225; Bouchard, 1978:37), and al- though many thorough crayfish surveys have been conducted in the south- eastern United States and neighboring areas it seems certain that additional species of primary burrowing crayfishes will be discovered here. Cambarus (Depressicambarus) deweesae, new species Figs. la-l Cambarus (Cambarus) diogenes.—Ortmann, 1931:154, 156, 157 [all in part]. Diagnosis.—Body and eyes with pigment. Rostrum without spines or tu- bercles. Areola 41.8 to 44.6 percent of total length of carapace (47.2 to 50.5 percent of postorbital carapace length) and obliterated or nearly so with space for no more than 1 punctation. Cervical spines reduced to small, 590 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON rounded tubercles; hepatic spines absent; branchiostegal spine present; sub- orbital angle lacking to obtuse; postorbital ridge moderately strong, rounded cephalically. Antennal scale narrow, longer than broad, broadest slightly distal to midlength. Chela with 2 rows of tubercles on mesial surface of palm, primary row with 6 to 8 tubercles (rarely 9), secondary row with 4 to 6 (rarely 7); opposable margin of dactyl with proximal 4 (rarely 3) tubercles prominent—first and fourth (rarely first and third) larger; corresponding margin of propodus with proximal 2 or 3 tubercles dominant in size, second or third largest; opposable margins of both with single row or irregular double row of denticles; lateral margin costate; lateral base of fixed finger impressed above, less so below. Hook on ischium of third pereiopod of male overreaching basioischial articulation and not opposed by tubercle on basis. First pleopod of first form male with central projection corneous, bladelike, with well-developed subapical notch, recurved at angle of approximately 100°; mesial process tumescent, tapering to 1 or 2 subacute tips and directed caudolaterally at angle of approximately 90° to shaft of appendage. First pleopod of second form male noncorneous; central projection rounded dis- tally; mesial process tapering to acute tip and longer than central projection. Annulus ventralis asymmetrical, subquadrangular, with caudal part some- what movable; cephalic half elevated and bearing longitudinal median trough between caudally divergent, longitudinal ridges; caudal half with sinuate sinus and elevated caudal wall. First pleopod of female uniramous and reaching approximately midlength of annulus ventralis when abdomen flexed. Holotypic male, form I.—Body subovate, vaulted (Fig. la). Abdomen narrower than thorax (13.7 and 19.9 mm). Greatest width of carapace greater than depth at caudodorsal margin of cervical groove (19.9 and 16.9 mm). Areola obliterated along part of its length and constituting 44.4 percent of total length of carapace (50.5 percent of postorbital carapace length) (Fig. 1k). Rostrum spatulate, deepened cephalically, with slightly convergent, thickened margins devoid of marginal spines or tubercles; upper surface with submarginal punctations and others scattered between. Acumen set off from proximal part of rostrum with concave, oblique margins and termi- nating in very small, upturned, corneous tubercle. Postorbital ridges mod- erately strong, deeply grooved dorsolaterally and rounded cephalically. Sub- orbital angle lacking; branchiostegal spine small (lacking on right side). Cervical spines represented by series of 3 or 4 rounded tubercles; hepatic area and lateral part of branchiostegites tuberculate, especially former; dor- sal part of carapace punctate. Abdomen shorter than carapace (32.4 and 41.4 mm); pleura short with caudoventral extremity broadly angular. Cephalic section of telson with single movable and immovable spines in each caudolateral corner, separated from caudal section by paired oblique excisions. Basal podomere of uropod VOLUME 92, NUMBER 3 591 Fig. 1. Cambarus (Depressicambarus) deweesae (a, b, c, d, e, h, j, kK, 1, holotype; i, allo- type; f, g, morphotype). a, Lateral view carapace; b, c, d, Lateral, caudal, mesial views first pleopod; e, Basis and ischium third pereiopod; f, g, Mesial and lateral views first pleopod; h, Antennal scale; i, Annulus ventralis; j, Epistome; k, 1, Dorsal view carapace and chela. 592 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON with very small spines extending over mesial and lateral rami. Lateral ramus of uropod with median ridge terminating in acute spine at transverse flexure; additional small ridge lateral to median one; proximal part with row of small spines distally and movable spine submarginally at caudolateral corner. Mesial ramus of uropod with median ridge terminating distally in premar- ginal acute spine. Caudal margin of tail fan with plumose setae; dorsal sur- face lightly setiferous. Cephalic lobe of epistome (Fig. 1j) pentagonal with slightly upturned ceph- alolateral margins and with small cephalomedian projection; ventral surface flattened. Basal part of epistome with deep, median fovea and pair of obliquely disposed, slitlike fossae immediately cephalic and subparallel to thickened, arched, short epistomal zygoma; lateral extremities without tu- bercles. Proximal segment of antennule with small spine on ventral surface at base of distal fourth. Antennae extending to fifth abdominal segment. Antennal scale (Fig. 1h) narrow, broadest slightly distal to midlength; thick- ened lateral part terminating in acute, corneous-tipped spine (broken on left) projecting forward but not reaching tip of rostrum; lamellar area narrow, with mesial margin angulate, crenulate and edged with long, plumose setae. Right chela (Fig. 11) approximately 2 times longer than broad (27.9 and 15.0 mm), well depressed, although slightly inflated proximolaterally; mesial margin of palm with 2 rows of 6 tubercles each in primary and secondary rows; dorsal surface bearing several small squamous tubercles over mesial half of palm. Lateral surface of propodus costate with row of punctations rendering proximolateral margin of fixed finger and palm irregular in dorsal aspect. Fixed finger with proximolateral base impressed dorsally, less so ventrally; dorsal and ventral surfaces with distinct submedian ridges flanked by setiferous punctations; opposable surface with row of 5 tubercles along proximal three-fifths of finger, third largest, decreasing in size proximally; additional small tubercle (absent due to injury on right) present on lower level at base of distal third, and double row of minute denticles extending proximally from corneous tip of finger to third tubercle from base, inter- rupted by fourth and fifth. Dorsal and ventral surfaces of dactyl with median longitudinal ridges flanked by setiferous punctations; opposable margin with row of 8 tubercles, first and fourth larger; mesial margin of dactyl tuber- culate along nearly proximal half and punctate along distal half; double row of minute denticles extending from corneous tip to fourth tubercle from base, interrupted by fifth through eighth. Carpus longer than broad with deep, oblique furrow dorsally; mesial sur- face with large procurved spine near midlength and with cluster of 5 smaller tubercles situated ventrad of it and additional single one proximally; disto- ventral margin with strong spiniform tubercle and smaller, proximally dis- posed one; podomere otherwise punctate. Dorsodistal surface of merus with 4 subacute tubercles and irregular row VOLUME 92, NUMBER 3 593 of 9 along crest of podomere; ventral surface with lateral row of 6 tubercles, some corneous-tipped, and mesial row of 12 corneous-tipped, acute tuber- cles, both rows decreasing in size proximally. Ischium with 2 very small tubercles on mesial margin. Hook on ischium of third pereiopod only (Fig. le); hook simple, over- reaching basioischial articulation and not opposed by tubercle on basis. Coxa of fourth pereiopod with prominent caudomesial boss; fifth pereiopod without prominence. For measurements see Table 1. First pleopods (Fig. 1b, c, d) reaching caudal part of coxae of third pe- reiopods when abdomen flexed. See ‘*Diagnosis’’ for description. Allotypic female.—Differing from holotype in following respects: subor- bital angle slightly more angulate. Cephalic part of epistome with more angular cephalolateral margins. Cervical spines represented by 2 or 3 dom- inant tubercles on each side. Chela with primary row of tubercles on mesial margin of palm consisting of 8 and 7 on right and left ones, respectively; secondary row of both with 5. Opposable margin of right propodus with row of 7 tubercles (8 on left), and single to double row of denticles extending proximally from corneous tip to fourth tubercle on left cheliped, interrupted by fifth through seventh tubercles (fourth through sixth on right). Opposable margin of dactyl with 10 and 12 tubercles on right and left chelipeds, re- spectively, fourth largest, and single row of denticles extending proximally from corneous tip to sixth tubercle on right cheliped (fifth on left), inter- rupted by seventh through tenth and sixth through eleventh tubercles on right and left fingers, respectively. Right carpus with cluster of 8 tubercles (7 on left) near large procurved spine on mesial margin. Merus with 9 tu- bercles along crest of podomere; ventral surface with 9 tubercles in lateral row and 13 in mesial one. Ischium with row of 4 small tubercles on mesial margin. Annulus ventralis (Fig. 11) subquadrangular, broader than long and situ- ated deep in sternum with cephalic part fused to sternum and caudal half somewhat movable. Annulus ventralis divided by sinus into C-shaped and triangular parts, latter with basal tongue projecting into concavity of “‘C.”’ Cephalic half elevated and bearing sinus broadening into median, longitu- dinal trough flanked by caudally diverging ridges; caudal portion elevated and nearly bisected by shallow sinus. Postannular sclerite approximately half as wide as annulus, with oval elevation (ventrally) almost one-third width of annulus. Morphotypic male, form II.—Differing from holotype in following re- spects: rostrum with margins less angulate cephalically; apical tubercle worn and barely visible. Cervical spines consisting of 2 dominant tubercles on right side, 1 on left; suborbital angle slightly more angulate; antennal scale moderately wide. Chela with primary row of 7 tubercles on mesial margin of right palm, secondary row of both with 4. Opposable margin of right 594 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Table 1. Measurements (mm) of Cambarus (Depressicambarus) deweesae. Holotypic Allotypic Morphotypic Male, Form I Female Male, Form II Carapace Height 16.9 18.2 KK Width 19.9 21.5 KK Total length of carapace 41.4 44.5 rar Postorbital carapace length 36.4 38.9 oe Areola Length 18.4 19.0 Pee Rostrum Width 5.4 2) yi) 4.6 Length S022 5.6** a pe Chela Length, mesial margin of palm 9.3 9.6 6.7 Width, palm 15.0 14.7 10.6 Length, lateral margin 21 2) 29.4 SEs Length, dactyl 18.3 al 14.4 * Areola obliterated. ** Measurement after Bouchard, 1973:103. *** Distal tubercle broken. **** Carapace pliant and wrinkled. propodus with row of 7 tubercles (6 on left), and row of denticles extending proximally from tip to sixth tubercle on right cheliped, interrupted by sev- enth (extending to fifth and interrupted by sixth on right). Opposable margin of dactyl with 11 tubercles on each cheliped, and single row of denticles extending proximally from corneous tip to fifth tubercle on right cheliped, interrupted by sixth through eleventh (interrupted by fifth through eleventh on left cheliped). Right carpus with cluster of 11 tubercles (13 on left) near large procurved spine on mesial margin. Merus with 9 and 12 tubercles along crest of right and left podomeres, respectively; ventral surface with 7 tu- bercles in lateral row and 13 in mesial one of both chelipeds. Hook on ischium of third pereiopod much reduced, not reaching basiois- chial articulation, and not opposing tubercle on basis; boss of coxa of fourth pereiopod somewhat smaller and less sharply defined. First pleopods (Fig. If, g) of uniform texture and reaching caudal portions of coxae of third pereiopods when abdomen flexed. See ‘‘Diagnosis’’ for description. Color notes.—Cephalothorax and abdomen concolorous, ranging from bright red or orange red to more somber brick red dorsally. For brevity range of red hues referred to as reddish. Gastric region and abdomen bright- VOLUME 92, NUMBER 3 595 er than hepatic area and former with paired, mottled areas marking attach- ment of mandibular muscles. Branchiostegites and hepatic areas concolor- ous reddish dorsally, fading to white ventrad. Pleural regions of abdomen reddish, cream or white ventrally and may bear paired light spots on first segment. Ventral aspects of cephalothorax and abdomen white. Rostral margins and postorbital ridges yellow or same color as cephalothorax. Tu- bercles, spines and articular condyles yellow or cream. Antennal scale red- dish, lateral margin darker than lamellar portion. Antennae reddish to brown. Chelae reddish with lighter proximolateral area; white ventrally. Distal ends of fingers yellow to cream or reddish. Pereiopods light reddish to pink dorsolaterally; white ventrolaterally. Distal podomeres darker dorsally than proximal ones. Ortmann (1931:157) noted the color of this species as follows: ‘‘. . . at the eastern foot of Walden Ridge, at Harriman and Dossett, all specimens found (2 ¢ II, 1 2) had a reddish ground color. While the female from Dossett has been recorded as completely ‘bright red,’ the males from Harriman are recorded as, ‘body olive-brown to reddish-brown (on carapace); rostrum and chelae orange-red; sides of carapace pale grayish-olive.’ ”’ Type-locality.—Seepage area near east bank of Poplar Creek at Tennessee State Highway 61 (south side of highway) in Anderson County, Tennessee. This locality lies within the Clinch River system (Tennessee River basin). Collected with C. deweesae from burrows at the type-locality was C. (Cam- barus) bartonii (Fabricius, 1798:407). Disposition of. types.—The holotypic male, form I (no. 148363) and the allotypic female (no. 148364) are deposited in the National Museum of Nat- ural History, Smithsonian Institution, Washington, D.C., U.S.A. The mor- photypic male, form II (no. 74.1092) is on deposit at the Carnegie Museum of Natural History, Pittsburgh, Pennsylvania, U.S.A. Paratypes consisting of 52 are in the Smithsonian Institution, 1¢ II and 12 in the Carnegie Mu- seum and 1¢1, 32, 1d juvenile, 12 juvenile and 1? bearing eggs are in the collection of the senior author. Range and specimens examined.—This crayfish is known from four lo- calities in a small area of Anderson and Roane counties, Tennessee, within the Clinch (Anderson County) and Emory (Roane County) River systems (Tennessee River basin). These localities lie near the foot of the eastern escarpment (Walden Ridge) of the Cumberland Mountains and Cumberland Plateau sections of the Appalachian Plateaus province in the Ridge and Valley province. Other primary burrowers that occur in the Ridge and Val- ley are C. JVugicambarus) dubius Faxon (1884:114), C. (Depressicambarus) striatus Hay (1902:437), C. (D.) cymatilis Hobbs (1970:241), C. (L.) di- ogenes (sens. lat.) and an undescribed Cambarus of the subgenus Lacuni- cambarus. Cambarus bartonii, regarded as a tertiary burrower (see Bou- 596 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON chard, 1976:588), has been collected from burrows some distance from surface waters, and is the only species collected with C. deweesae to date. TENNESSEE—ANDERSON COUNTY—(1) Type-locality. [1V/20/71. D. A. Etnier and J. P. Dewees, coll. 5°; (ii) Type-locality. II/13-16/72. D. A. E. and F. L. Oakberg, coll. 2d 1, 22, 12j, 1 ovigerous @; (iii) Swamp, Dossett (foot of Walden Ridge). IX/2/14. A. E. Ortmann, coll. 12. ROANE CouUN- TY—(iv) Field between County Road 2437 and Tennessee State Highway 61, west of the town of Little Emory. I/25/71. D. A. E., coll. 12, 14); (v) Harriman (foot of Walden Ridge). V/16/15. A. E. O., coll. 2d II. Variations.—The small number of specimens available exhibit little vari- ation other than the usual expected range in meristics and abrasion of spines and tubercles in late intermolt stages. Size.—The largest specimen available is a female with a carapace length of 48.0 mm (postorbital carapace length 42.0 mm). The smallest first form male has corresponding lengths of 41.3 and 36.8 mm. The largest first form male has a carapace length of 41.4 mm (postorbital carapace length 36.4 mm). The only female specimen collected with eggs has a carapace length of 46.4 mm (postorbital carapace length 41.3 mm). Life history notes. —Form I males were collected in March 1972 between the thirteenth and sixteenth of the month (exact date unknown), and a single ovigerous female was collected during this same period. Ecological notes. —Cambarus (D.) deweesae is a primary burrower. A label accompanying the crayfish collected by Ortmann from Dossett indi- cates this individual to have been collected from a swamp. All of Ortmann’s specimens were ‘‘dug out of holes’’ (1931:155). The type-locality is a poorly drained floodplain on the east side of Poplar Creek and on the south side of Tennessee State Highway 61, Anderson County, Tennessee. Soils consist of clays interspersed with fragments of shale. Burrows are complex and variable, with much of this variability attributable to obstructions presented by large shale fragments. Typical bur- rows consist of a conspicuous opening surrounded by excavated clay. Well- defined chimneys have not been observed. The shaft leading from this open- ing is vertical or nearly so. One to several short, horizontal culs-de-sac often extend from the vertical shaft approximately 5 to 20 cm below the soil surface. At a depth of 50 to 70 cm the vertical shaft begins to assume a more horizontal position. Often the burrow bifurcates at this depth, with one of the resulting more horizontal galleries a cul-de-sac and the other angling back toward the surface. The opening of the latter branch is well concealed under vegetation, may have a short, horizontal, blind chamber near the surface and lacks excavated material around the opening. Digging the entire burrow system is a tedious but effective method of collecting specimens. Often individuals can be obtained by using a plunging motion with an ex- tended forearm. Displaced water can be heard gurgling to the surface at the site of the concealed opening, and the resident of the burrow may be dis- Vy aa my Fig. 2. Cambarus (Depressicambarus) striatus, lateral views of left first pleopod, male, form I: a-b, Winston Co., MS; c, Dawson Co., GA; d, Montgomery Co., TN; e, Loudon Co., TN; f, Coffee Co., TN; g, Blount Co., AL; h, DeKalb Co., AL; i, Marion Co., AL; j, Lafayette Co., MS; k, Bledsoe Co., TN; 1, Henderson Co., TN; m, Tuscaloosa Co., AL; n, Macon Co., AL; 0, Lee Co., AL. 598 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON placed to the mouth of the concealed opening. A burrow is typically occu- pied by a single animal, but the high population density occasionally results in adjacent burrows being connected by heavily silted, horizontal tunnels that are presumably little used. Although considerable construction and filling has recently taken place near the type-locality, this crayfish was still thriving in March 1978. Much of the area will become incorporated into a golf course that is currently under construction. Since burrowing crayfish are typically compatible with most areas of golf courses, the continued survival of this species at the type- locality seems certain. Relationships. —Cambarus deweesae is a member of the subgenus De- pressicambarus, exhibiting a broadly subtriangular and depressed chela with two major rows of tubercles along the mesial margin of the palm, a common feature of the group. Within this subgenus C. deweesae bears closest affin- ities to members of the latimanus group (Bouchard, 1978:44) and in partic- ular C. striatus Hay, a wide-ranging and variable species adapted to a broad array of habitats. Cambarus striatus occupies epigean waters as well as burrows as a primary, secondary or tertiary burrower (Bouchard, 1976:591, slightly modified from Hobbs, 1942:20). The most distinctive differing features between the two species are the first pleopod of the first form male and the annulus ventralis of the female. In C. deweesae the central projection of the gonopod is short, tapered and bears a subapical notch, a combination of features not known from any population of C. striatus (cf. Fig. 1b and Fig. 2). In the female of C. de- weesae the cephalic part of the annulus ventralis is elevated while in C. striatus the cephalic part is distinctly lower in height than the caudal wall. An elevated cephalic part of the annulus ventralis is also found in members of the halli group of the subgenus and in particular in C. obstipus Hall and populations of C. englishi Hobbs which occur syntopically with C. halli Hobbs. ) The areola of C. deweesae is obliterated or very narrow with space for no more than one punctation while in C. striatus the areola is rarely oblit- erated and usually slightly wider. The suborbital angle in C. deweesae is lacking or poorly developed, while it is generally moderately well developed in C. striatus. The reddish color pattern of C. deweesae is unknown in populations of C. striatus, a species which basically exhibits a color pattern consisting of browns and greens, although occasional individuals with blue or blue-gray colors have been observed (see Bouchard 1978:42). Within the subgenus Depressicambarus the reddish color of C. deweesae also occurs in C. pyronotus Bouchard (orange-red, Bouchard, 1978:39) and has been observed rarely in C. graysoni (op. cit.). Cambarus pyronotus like C. deweesae exhibits a reduced suborbital angle (obsolete in C. pyronotus). Both species occur at the periphery of the range of C. striatus—C. pyro- 4 VOLUME 92, NUMBER 3 599 notus in Torreya Ravine, Liberty County, Florida, and C. deweesae in Anderson and Roane counties, Tennessee, near the foot of Walden Ridge. Joel Palmer Dewees 1943-1977 Etymology.—We are pleased to name this new species of crayfish in honor of the late Joel P. Dewees (1943-1977), whose interest in burrowing cray- fishes has enriched our knowledge of many species in the Appalachian Pla- teaus province and southern parts of the Ridge and Valley and Blue Ridge provinces. Miss Dewees also assisted in the collection of material from the type-locality. Acknowledgments.—We should like to thank Dr. Horton H. Hobbs, Jr., Smithsonian Institution, and Judith W. Bouchard for kindly reviewing the manuscript and the latter for rendering the figures. To Dr. J. J. Parodiz, Carnegie Museum of Natural History, we are indebted for the loan of ad- ditional specimens of the species described herein. The senior author wishes to thank the Smithsonian Institution for awarding him a Smithsonian Post- doctoral Fellowship which permitted a study of the subgenus Depressicam- barus and the recognition of this new species. In addition, a research grant, USDI 14-16-0008-2010, from the Office of Endangered Species, United States Department of the Interior, is gratefully acknowledged. 600 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Literature Cited Bouchard, Raymond W. 1973. A new crayfish of the subgenus Jugicambarus from Tennessee with an emended definition of the subgenus (Astacidae, Decapoda).—American Midl. Natur. 89(1):103-111. . 1976. Geography and ecology of crayfishes of the Cumberland Plateau and Cumber- land Mountains, Kentucky, Virginia, Tennessee, Georgia and Alabama. Part II. The genera Fallicambarus and Cambarus.—Pp. 585-605 in James W. Avault, Jr., ed. Fresh- water Crayfish. Papers from the Second International Symposium on Freshwater Cray- fish, Baton Rouge, Louisiana, U.S.A. Division of Continuing Education, Louisiana State University. . 1978. Taxonomy, ecology and phylogeny of the subgenus Depressicambarus with the description of a new species from Florida and redescriptions of Cambarus graysoni, Cambarus latimanus and Cambarus striatus (Decapoda: Cambaridae).—Bull. Alabama Mus. Natur. Hist. 3:27—60. Fabricius, Johann C. 1798. Supplementum Entomologiae Systematicae.—Hafniae:Proft et Storch. 572 pp. Faxon, Walter. 1884. Descriptions of new species of Cambarus, to which is added a syn- onymical list of the known species of Cambarus and Astacus.—Proc. Amer. Acad. Arts and Sci. 20: 107-158. Girard, Charles. 1852. A revision of the North American astaci, with observations on their habits and geographical distribution.—Proc. Acad. Natur. Sci. Philadelphia 6:87—91. Hay, William P. 1902. Observations on the crustacean fauna of Nickajack Cave, Tennessee, and vicinity.—Proc. U.S. Nat. Mus. 25(1292):417-439. Hobbs, Horton H., Jr. 1942. The crayfishes of Florida.—Univ. Florida Publ., Biol. Sci. Ser. 3(2):v + 179 pp. . 1967. The current status of the crayfishes listed by Girard (1852) in his “‘A Revision of the North American Astaci... .”—Crustaceana 12(2): 124-132. . 1970. New crayfishes of the genus Cambarus from Tennessee and Georgia (Decapoda, Astacidae).—Proc. Biol. Soc. Washington 83(24):329-348. . 1973. New species and relationships of the members of the genus Fallicambarus.— Proc. Biol. Soc. Washington 86(40):461—482. . 1975. New crayfishes (Decapoda: Cambaridae) from the southern United States and Mexico.—Smithsonian Contrib. Zool. 201:iii + 34 pp. Ortmann, Arnold E. 1931. Crawfishes of the Southern Appalachians and the Cumberland Plateau.—Annals Carnegie Mus. 20(2):61—160. Rafinesque, Constantine S. 1817. Synopsis of four new genera and ten new species of Crus- tacea, found in the United States —American Monthly Mag. and Critical Rev. 2:40—43. Schuster, Guenter A. 1976. A new primary burrowing crayfish of the subgenus Jugicambarus (Decapoda, Cambaridae) from Kentucky, with notes on its life history —American Midl. Nat. 95(1):225-—230. (RWB) 7500 Seaview Avenue, Wildwood Crest, New Jersey 08260; (DAE) Department of Zoology, University of Tennessee, Knoxville 37916. PROC. BIOL. SOC. WASH. 92(3), 1979, pp. 601-605 A NEW NORTH AMERICAN SPECIES OF FRESH-WATER TUBIFICIDAE (OLIGOCHAETA) Michael S. Loden Abstract.—Sexually mature specimens of Tubifex harmani n. sp. were collected from Louisiana, although immature forms apparently of this species have been recorded from Indiana, Florida, and British Columbia. The male genital system shows affinities to Tubifex, Isochaeta, and Ilyod- rilus. Asexual reproduction by fragmentation is characteristic. Specimens of a previously undescribed tubificid, Tubifex harmani n. sp., were collected from among the roots of Colocasia antiquorum (L.) Schott (Fam. Araceae) from a lake adjacent to the Baton Rouge campus of the Louisiana State University. The particulars of the male reproductive tract, determined from stained, whole-mount specimens, show affinities to the genera Tubifex Lamarck, Isochaeta Pointner, and Ilyodrilus Eisen. Tubifex harmani new species Fig. 1A—E Material examined.—HOLOTYPE: NMNH 56184, Louisiana: East Baton Rouge Parish; east side of University Lake, Baton Rouge; 19 October 1976; M. S. Loden. PARATYPES: NMNH 56185, same locality and date, 4 speci- mens; LSU 2051, same locality and date, 3 specimens. ADDITIONAL SPEC- IMENS: Florida: Jefferson Co., ditch along US 98, 0.5 mi E Wakulla Co. line, 8 March 1978, W. J. Harman and M. S. Loden. LSU 2052, 1 specimen. Indiana: Gibson Co., Public Service Indiana Pump Storage Reservoir, 3 April 1975, LSU 2053, 2 specimens. British Columbia: Victoria, Victoria Airport, 5 May 1976, R. O. Brinkhurst Collection, 4 specimens. Etymology.—Named in honor of Professor Walter J. Harman of Louisiana State University. Description.—Length 10-30 mm (preserved). 50-95 segments. Diameter 0.2 mm at segment XI, 0.15 mm at posterior end. Prostomium rounded to triangular, as long as width at peristomial junction. Clitellum between in- tersegmental grooves 9/10 and 12/13. Anterior dorsal setal bundles contain- ing 1-3 (usually 2) hair setae and 2-3 pectinate crotchet setae. Hairs very finely serrated, up to 500 um long. Pectinate setae 63-91 um long, 3 wm diameter; lateral teeth subequal, distal thinner; 1—4 fine intermediate teeth. Dorsal bundles in clitellar and immediately postclitellar segments with 1-2 (usually 1) hairs up to 425 wm long, and 2 crotchets 70-85 um long, 4—4.5 602 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Cc D E Fig. 1. Tubifex harmani A, Male duct; B, Penis sheath; C, Pectinate seta; D, Anterior ventral seta; E, Posterior crotchet seta. Based on Zeichentubus drawings of stained, whole- mount specimens. ym thick; distal tooth thinner and straighter, proximal tooth slightly re- curved, occasionally with one thin intermediate tooth. Posterior dorsal bun- dles with 1-2 hairs, 265-300 um long, and 1-2 bifid crotchets; crotchets strongly sigmoid, 70-90 .m long, 9-12 wm thick distally; distal tooth shorter and thinner than stout, recurved proximal; nodulus distal. Anterior ventral setal bifid, 3-S per bundle; in II 49-58 ym long, up to 10 um thick, distal tooth longer and thinner, nodulus median. In III—-X ventrals 60-81 um long, up to 10 wm thick; teeth subequal to distal slightly shorter; distal tooth thinner. Posterior ventrals resemble posterior dorsal crotchets, occasionally slightly longer and thinner. Male genital system with all structures paired. Slightly conical male fun- VOLUME 92, NUMBER 3 603 nels ca. 90 ~m maximum diameter on septum 10/11. Vasa deferentia ap- proximately 600 um long, 25 wm diameter, ciliated anteriorly from septum 11/12. Atria in XIII, 380 wm long, 30-58 wm wide, surrounded by muscle fibers; joined subapically and posteriorly by compact prostate glands, 420 wm long, 120 wm maximum width. Ejaculatory ducts in XI, 250 um long, 30-53 wm wide, transitioning into penes. Thin cuticular penis sheath 58 um long, basal diameter 20 um; head at an angle, oval, maximum diameter 28 ym. Male pores ventrolateral in XI, in transverse setal plane, just lateral to the setal bundles. Spermathecae paired in segment X. Ampullae ovoid, 163 wm long, 117 ym wide; duct 35 wm diameter, 200 um long. Spermatozeugmata ca. 120 wm long, widened at one end; diameter 51 wm at wide end, 23 um at narrow end; axial cavity 9-25 wm diameter. Spermathecal pores ventrolateral in X, in transverse setal plane, just lateral to the setal bundles. No modified genital setae. Asexual reproduction by fragmentation, 10 segments formed anteriorly. Discussion.—The uniquity of this species in the Tubificinae is based on the morphology of the setae, the male genitalia, and on evidence of asexual reproduction. The posterior setae resemble those of several species of the Peloscolex complex and Psammoryctides, but the configuration of the male ducts precludes conspecificity of T. harmani with any in those genera. Suggestions of asexual reproduction first came from the discovery of sev- eral specimens that showed anterior regeneration from segments with setae characteristic of the posterior third of the body; the crotchet setae of seg- ment XI were of the strongly sigmoid type. In one individual the periproct was separated from the zone of anterior regeneration by only 35 segments. The suggestions of asexual reproduction were reinforced when specimens in cultures were severed approximately one-third of the body distance from the anterior (ca. segment XXX). Both anterior and posterior portions de- veloped into complete worms in all cases, thus the regenerative capacity of the species is great. Two worms autotomized and regenerated spontaneously. The type-locality was sampled systematically for 18 months, and occa- sionally for an additional 12 months; of well over 300 specimens collected, only three individuals were sexually mature. Asexual reproduction, there- fore, appears to be a primary mode of propagation. No sexual maturity was observed in a population of a species of Bothrioneurum at the same location; this genus has species that characteristically reproduce asexually (Brink- hurst, (1971). Although sexually mature worms are known only from the type-locality, immature specimens possessing setae virtually identical to those described for T. harmani have been collected from Florida, Indiana, and British Co- lumbia; these specimens have been identified tentatively as T. harmani. Tubifex harmani presents features that may cause it to be placed by some 604 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON investigators in [sochaeta (sensu Brinkhurst, 1963) or Ilyodrilus. Brinkhurst stated in his definitions that Isochaeta ‘‘has elongate, narrow atria scarcely distinguishable from vasa deferentia’’; Tubifex was defined as having ‘‘vasa deferentia at least as long as the pear-shaped to cylindrical atria which are frequently turned over apically, otherwise standing vertically in XI; no dis- tinct ejaculatory ducts; prostate glands large, closely connected to the atria subapically on anterior side.’’ Ilyodrilus, in turn, has “‘large prostate glands on the upper posterior walls, ejaculatory ducts separated from atria, but broad.”’ The atria of T. harmani are elongate and rather narrow; the prostate connections are subapical and posterior. Only the presence of muscle fibers distinguishes atria from ejaculatory ducts. The vasa deferentia are conspic- uous by their reduced diameter, and they are ciliated in the proximal por- tion. Thus, characters of each genus are present, but Tubifex and Isochaeta appear to have more in common with the new species. The genus [sochaeta was erected by Pointner (1911) for I. virulenta. The type-material was examined by Brinkhurst (1963), who supported Pointner’s views on the validity of the species. The types were also examined by Hrabé (1966), who agreed with Michaelsen (1926) that J. virulenta was based on specimens that were, in actuality, incompletely mature individuals of a Lim- nodrilus species, probably L. udekemianus Claparede. Hrabé therefore con- cluded that not only was the species invalid, but that the genus was a junior synonym of Limnodrilus. Brinkhurst (1963, 1971) included species with and without hair setae in both Isochaeta and Tubifex. Hrabé (1966, 1967) placed in Tubifex only species with hair setae (except some subspecies of T. tubifex); he created the genus [sochaetides for accomodating similar species that lack hair setae. Thus, there is the situation where two of the leading authorities in the field differ in opinion not only on the validity of the genera, but on the relative importance of the details of the genitalia and setal configurations. If Hrabé’s interpretation of tubificine genera is accepted, T. harmani clearly would fall into Tubifex. With Brinkhurst’s interpretation, the species could be placed into either Tubifex or Isochaeta, depending upon the rel- ative weights ascribed to the various genital characters. Dr. R. O. Brinkhurst (pers. com.) agreed with the uncertainty of the generic placement of T. harmani, but he was more favorably disposed to- ward Isochaeta. With all due respect to Dr. Brinkhurst, I have elected to place the new species in Tubifex due to: 1. the large prostate being more typical of Tubifex. 2. the vasa deferentia being clearly distinguishable from the atria. 3. the ability to distinguish the atrium from the ejaculatory duct only by the presence of atrial muscle fibers. VOLUME 92, NUMBER 3 605 4. the ease of relegating the species to Hrabé’s (1966) definition of Tubifex. 5. my subjectivity in assigning the species to the older genus while there is uncertainty in the generic placement. Species of Isochaeta and Tubifex (sensu Brinkhurst, 1963, 1971) that share with T. harmani the presence of hair setae and a well-defined penis sheath, and an absence of genital setae include J. nevadana Brinkhurst, J. israelis Brinkhurst, T. ignotus (Stol¢), T. montanus Kowalewski, T. kessleri Hrabé, and TJ. nerthus Michaelsen. In addition to differing from 7. harmani in the configurations of the male ducts, none of those species have the enlarged bifid setae in the posterior dorsal and ventral bundles. Literature Cited Brinkhurst, R. O. 1963. Taxonomical studies on the Tubificidae (Annelida, Oligochaeta).— Int. Rev. Ges. Hydrobiol. (Syst.) 2:7-89. . 1971. Family Tubificidae.—Pp. 444-625 in R. O. Brinkhurst and B. G. M. Jamieson, Aquatic Oligochaeta of the World. Hrabé, S. 1966. New or insufficiently known species of the family Tubificidae.—Publ. Fac. Sci. Purkyné Univ. No. 450:101-112. . 1967. Two new species of the family Tubificidae from the Black Sea, with remarks about various species of the subfamily Tubificinae.—Publ. Fac. Sci. Purkyné Univ. No. 485:331-356. Michaelsen, W. 1926. Oligochaten aus dem Ryck bei Greifswald und von benachbarten Meers- gebieten.—Mitt. Zool. Inst. Hamburg 42:21-29. Pointner, H. 1911. Beitrage zur Kenntnis der Oligochatenfauna der Gewasser von Graz.— Zeitschr. Wiss. Zool. 98:626—676. Department of Zoology, Louisiana State University, Baton Rouge, Lou- isiana 70803. PROC. BIOL. SOC. WASH. 92(3), 1979, pp. 606-617 REVISION OF SOME POLYDORIDS (POLYCHAETA: SPIONIDAE) DESCRIBED AND RECORDED FROM BRITISH COLUMBIA BY EDITH AND CYRIL BERKELEY James A. Blake Abstract.—An examination of some spionid polychaetes from the collec- tions of the late Edith and Cyril Berkeley from British Columbia resulted in the following revisions: (1) Polydora caeca var. magna E. Berkeley, 1927, P. magna: Berkeley and Berkeley (1936), P. socialis plena Berkeley and Berkeley, 1936 and P. caeca: Berkeley and Berkeley (1936) are syn- onymized with P. socialis (Schmarda, 1861); (2) P. cardalia E. Berkeley, 1927 is redescribed; (3) P. (Boccardia) natrix (sensu Berkeley and Berkeley, 1936) is redescribed as Boccardia pugettensis n. sp. The different forms of specialized posterior notosetae, found among some species of the polydorid complex, are discussed. The late Edith and Cyril Berkeley, either singly or jointly, published more than 50 papers dealing with the polychaete fauna of western North America (Pettibone, 1967:19-21; Arai, 1971). Their 2 faunal treatments (Berkeley and Berkeley, 1948; 1952) were the major source of information for the identi- fication of Canadian Pacific polychaetes for many years. The bulk of their collections, including most of the types, are now lodged in the Smithsonian Institution (Pettibone, 1967). The Berkeleys described or recorded approx- imately 25 species of spionid polychaetes from British Columbian waters. Several species have been revised or commented upon by Pettibone (1962), Woodwick (1963a; b), Blake (1966; 1971), Foster (1971), Blake and Wood- wick (1972) and Banse (1972). The following spionid species from British Columbia are included in this report: Polydora socialis (Schmarda, 1861). Includes: P.caeca var. magna E. Berkeley, 1927. P. magna: Berkeley and Berkeley (1936). P. socialis plena Berkeley and Berkeley, 1936. P. caeca: Berkeley and Berkeley (1936). [Not Oersted, 1843.] Polydora cardalia E. Berkeley, 1927. Boccardia pugettensis, new species. Includes: P. (Boccardia) natrix: Berkeley and Berkeley (1936). [Not Soderstrom, 1920.] VOLUME 92, NUMBER 3 607 Specimens from the Smithsonian Institution (USNM), including the Berkeley collection, were loaned by Marian H. Pettibone and Meredith L. Jones. Additional comparative materials were provided by the late Kathar- ine D. Hobson, British Columbia Provincial Museum (BCPM), Kristian Fau- chald and Karen Green of the Allan Hancock Foundation (AHF), Dusty Chivers, California Academy of Sciences (CAS) and Lars Orrhage, Swedish Museum of Natural History, Stockholm (SMNH). I am grateful to all of the above-named individuals for their assistance and to William J. Light for reading and commenting on this paper. Polydora socialis (Schmarda, 1861) Leucodore socialis Schmarda, 1861:64, figs. a—c, pl. 26:fig. 209. Polydora caeca var. magna E. Berkeley, 1927:419.—Pettibone, 1967:11 [New Synonymy]. Polydora magna: Berkeley and Berkeley, 1936:473; 1952:21 [New Synon- ymy]. Polydora socialis plena Berkeley and Berkeley, 1936:468—469; 1952:22.— Pettibone, 1967:11.: Fide Biake, 1971. Polydora caeca: Berkeley and Berkeley, 1936:469; 1952:20-21, not figs. 36— 37, taken from Fauvel, 1927. [Not Oersted, 1843 sensu Fauvel, 1927 and authors. ] Polydora socialis: Hartman, 1941:310-311, pl. 48, figs. 41-42; 1969:147.— Blake, 1971:20—23, figs. 13-14 [Synonymy]; 1975:215, figs. 237-238.— Light, 1977:71; 1978:179-181, fig. 180a—l—Blake and Kudenov, 1978: 248-250, fig. 38d—e. Polydora plena: Foster, 1971:24—25, figs. 22-29. Fide Light, 1977. Material examined.—CANADA: British Columbia, Departure Bay, Na- naimo District, clean sand beach, littoral, 1 May 1935, coll. E. and C. Berke- ley, holotype (USNM 32705) and 22 paratypes (USNM 32704) of Polydora socialis plena. Nanaimo, low tide, coll. E. and C. Berkeley, holotype of Polydora caeca magna (USNM 32710). West coast of Vancouver Island, 137 m, 15 May 1934, coll. E. and C. Berkeley, from walls of Nicomache lumbricalis tubes (Maldanidae), 7 specimens identified as Polydora caeca (USNM 36737). Cordova Bay, Vancouver Island, intertidal, sand, 27 July 1973, coll. B. Turner, 5 specimens (BCPM 973.151.4N).—CALIFORNIA: San Francisco Bay, San Bruno, seaplane harbor channel marker “‘1,’’ 4 m, coll. F. Nichols, 10 specimens (CAS 000779). Numerous additional speci- mens from Tomales Bay and Morro Bay, author’s collections. Remarks.—Polydora caeca magna E. Berkeley and P. caeca (sensu Berkeley and Berkeley, 1936) are here referred to P. socialis. The synon- ymy of P. socialis plena Berkeley and Berkeley, 1936 with P. socialis sensu stricto proposed by Blake (1971) is reconfirmed. The specimens examined 608 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON agree in all significant details with the descriptions of P. socialis presented by Blake (1971) and Light (1978). The holotype of P. caeca magna is about 100 mm long and its caruncle extends posteriorly for 12 setigers. The great body length and long caruncle were cited as the primary characters distin- guishing this species from P. caeca (E. Berkeley, 1927; Berkeley and Berke- ley, 1936). Caruncle length, however, appears to be size-dependent in P. socialis. The caruncle lengths of New England specimens of P. socialis vary from 5-9 setigers, with those specimens having the longest caruncles also having a greater body length, up to 55 mm (Blake, 1971; also unpublished data). It is likely that specimens having even greater body lengths, such as seen in the holotype of P. caeca magna, would also have longer caruncles. Berkeley and Berkeley (1936; 1952) noted that their specimens identified as P. caeca had awl-shaped dorsal setae in posterior notopodia, while the types of P. magna had very sharp, slender capillaries with narrow wings. I have examined the posterior notopodial setae of the Berkeleys’ specimens and others from British Columbia very carefully and in no instance have awl-shaped spines (these usually considered to be acicular) been observed. Instead, most posterior notopodia bear 7-8 long and 5-6 short, nearly smooth capillary setae, with occasional notopodia lacking most or all of the short capillaries. The short capillaries, rather than being acicular spines, are of the same thickness as the longer capillaries and appear to represent mere- ly stages in the emergence, growth and replacement of the longer capillaries. These short capillaries are called ‘“‘needlelike’’ by Light (1978), but an ex- amination of his San Francisco Bay specimens indicates that they also ap- pear to be replacement setae for the longer capillaries. Thick acicular spines have been noted in the posterior notopodia of specimens of P. caeca from Europe (Blake, unpublished data). In those specimens, however, the spines are thicker than the accompanying capillaries. In P. socialis, the major spines of setiger 5 are generally straight, not falcate, and have a weakly developed subterminal boss or enlargement. These spines are similar in specimens from British Columbia, California and New England. The hooded hooks lack a manubrium and have a reduced angle between the 2 teeth and a large angle between the main fang and shaft. A reduction of the apical tooth occurs in posterior neuropodia in some New England specimens (Blake, 1971) and in some San Francisco Bay specimens (Light, 1978; Blake, personal observation), but was not observed on the specimens from British Columbia. This variability does not appear to be taxonomically significant. The pygidia of British Columbian specimens are very similar to those figured by Blake (1971) in having a large ventral lobe and 2 small dorsal ones. A gizzard-like structure (Blake, 1971) occurs at the level of setigers 16-18 in all specimens examined. Polydora socialis is a widespread and probably cosmopolitan species. The VOLUME 92, NUMBER 3 609 small differences observed between widely separated populations do not exceed the expected genetic variability for such eurytopic forms. Distribution.—East and west coasts of North America; Gulf of Mexico; Chile; Falkland Islands; Australia. Polydora cardalia E. Berkeley, 1927 Figs. 1, 2 Polydora cardalia E. Berkeley, 1927:418—419, pl. I, fig. 14.—Berkeley and Berkeley, 1952:21, figs. 38—39.—Pettibone, 1967:11. Material examined.—CANADA: British Columbia, Nanaimo, Cardale Point, 6 July 1921, dredged off Round Island, 19 Aug. 1920, False Narrows, 20 May 1920, Rocky Bay, 16 May 1920, 21 paratypes (USNM 32708). Description.—A large species, up to 35 mm long and 1.5 mm wide for about 160 segments; reported up to 140 mm long and 2 mm wide for 400 segments (Berkeley, 1927). Body dark brown in alcohol, with small trans- verse pigment bands on some anterior segments. Prostomium deeply incised on anterior margin, forming 2 divergent lobes (Fig. 1A); caruncle extending posteriorly to middle or end of setiger 5; no occipital tentacle; 2 pairs of eyes, anterior pair cup-shaped and laterally positioned, posterior pair oval and medially positioned. Peristomium broad; palps long, reaching posteriorly for about 20 setigers. Setiger 1 well-developed; notopodium with small fascicle of capillary no- tosetae and broad, leaflike postsetal lamella (Fig. 1A); neuropodium with fascicle of long capillary neurosetae and broad, elliptical postsetal lamella. Setigers 2, 3, 4, 6 and subsequent setigers with large spreading fascicles of unilimbate capillary notosetae arranged in 2 tiers, those of anterior tier being shorter than those of posterior tier; with broad notopodial postsetal lamellae on setigers 2—4 (Fig. 1A); notopodia in posterior region with 5-7 long, slen- der capillaries and 13-18 thin, needlelike capillaries (Fig. 1E). Neuropodia of setigers 2, 3, 4 and 6 with capillaries similar to those of notopodia; be- ginning on setiger 7, neuropodium with 5—6 bidentate hooded hooks accom- panied by 2-3 slender capillaries in superior position; hooks with reduced angle between teeth and wide angle between main fang and shaft, without constriction on shaft, and with fine serrations on margin of hood opening (Fig. 1B). Modified setiger 5 with dorsal fascicle of numerous, fimbriated geniculate setae, a curved row of 12 or more major spines alternating with bilimbate companion setae (Fig. 1C, D, 2), and a ventral fascicle of unilimbate capil- laries; major spines simple, lacking accessory structures, subterminal en- largement poorly developed, but each spine with numerous fine, scratch- like etchings in subterminal location when viewed at 1000 (Fig. 2). 610 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Gy, gis: ek a Yj “ e . / a BI a8 RA Fig. 1. Polydora cardalia (paratype, USNM 32708): A, Anterior end in dorsal view, palps missing; B, Hooded hook; C, Companion seta from setiger 5; D, Major spine from setiger 5; E, Posterior notopodium in dorsal view; F, Posterior end and pygidium in dorsal view. Branchiae from setiger 8, continuing to near posterior end. Pygidium 3- lobed in paratypes, but reported as 3- or 4-lobed by Berkeley (1927). Giz- zard-like structure present at level of setigers 14-16. Remarks.—Polydora cardalia appears to be transitional between P. so- cialis and P. flava in the development of fine needlelike posterior notosetae. In P. socialis, short capillaries are few in number and are believed to rep- resent developing replacement setae for the longer capillaries (see above). In P. cardalia, there are up to 18 thin capillaries projecting up to one-fourth of the distance from the notopodium to the tips of the longer capillaries; they are consistent on each segment and do not appear to be replacement setae. In P. flava, the needles form dense packets barely protruding above the cuticle (Fauvel, 1927; Blake and Kudenov, 1978). The subterminal en- largements of the major spines on setiger 5 of P. cardalia are reduced, but VOLUME 92, NUMBER 3 611 Fig. 2. Polydora cardalia (paratype, USNM 32708): Fascicle of major spines and compan- ion setae from setiger 5, with inset [not to scale] showing form of subterminal etchings on spines. the fine, scratchlike, subterminal etchings are unusual. Such markings have not previously been reported from the major spines of polydorids. Polydora socialis has 3—7 emergent spines on setiger 5 (Light, 1978; Blake, 1971, Fig. 14b), P. flava has S—6 such spines (Mesnil, 1896) and P. cardalia has 12 or more. P. cardalia is further unusual in having exceptionally large, flattened, glandular postsetal lamellae in the anterior notopodia. These lamellae on setiger | are so large that they overlap nearly one-half of the length of setiger 2. In P. socialis, the same structures are much smaller, more elliptical, and although often flattened, are never glandular and do not extend beyond the limits of setiger 1. Polydora cardalia may occur off southern California. Two specimens 612 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON from the BLM Outer Continental Shelf Survey (334-337 m), deposited in the Allan Hancock Foundation, were examined. Both are posteriorly incom- plete, but agree in other respects with P. cardalia, including the form of the anterior notopodia. Unfortunately, the type-specimens of P. neocardalia Hartman, 1961 could not be located among the collections of the Allan Hancock Foundation, precluding any determination of its status. Light (1978) suggested that P. neocardalia might be the same as P. socialis, but until the types can be located and studied, the status of P. neocardalia and the BLM specimens remains uncertain. Distribution.—British Columbia. Boccardia pugettensis, new species Fig. 3 Polydora (Boccardia) natrix: Berkeley and Berkeley, 1936:472; 1952:18, not Fig. 28, taken from Soderstrom (1920).—Banse, Hobson and Nichols, 1968:538. [Not Soderstrom, 1920. ] Material examined.—CANADA: British Columbia, Mudge Island, 16 June 1927, coll. E. and C. Berkeley, holotype (USNM 57479) and 2 para- types (USNM 39567). Clayoquot, 4 June 1946, coll. E. F. Ricketts, sand flats, | paratype (USNM 39565). Nanoose Bay, 6 May 1965, coll. D. Ellis, 30 m, sand-silt, 5 paratypes (BCPM 42).—WASHINGTON: Puget Sound, 47°44'31"N; 122°32'41"W, Sta. 4, 29 April 1963, coll. U. Lie, 12 m, 3 para- types (USNM 37017-9). Ballard, Seattle, 29 June 1946, coll. M. H. Petti- bone, 3 paratypes (USNM 45193). Description.—A large species, up to 25 mm long and 2.5 mm wide for 65 setigers. Color: light tan to dark brown in alcohol, with some brown pigment on prostomium and anterior setigers; small paired black spots on dorsum of some specimens. Prostomium incised, forming 2 distinct lobes, with caruncle continuing to posterior margin of setiger 2 (Fig. 3A); no occipital tentacle; 2 pairs of eyes, anterior pair more widely spaced and cup-shaped; posterior pair oval. Palps thick, extending posteriorly for 8—9 setigers. Setiger 1 smaller than subsequent setigers, but with well-developed noto- and neuropodia, each bearing fascicles of capillary setae. Notopodia of se- tigers 2, 3, 4, 6 and subsequent setigers with prominent fascicles of unilim- bate capillaries arranged in 3 tiers: first tier with short, thick setae, second, with setae gradually elongating, third, with setae longest and thinnest; no- topodia of far posterior setigers with 4—5 long capillaries and 2-3 short, pointed spines (Fig. 31, J). Neurosetae of setigers 2, 3, 4 and 6 in 3-tiered fascicles similar to notosetae; beginning on setiger 7, neuropodium with S— 6 bidentate hooded hooks accompanied by 7-8 inferior unilimbate capillar- ies, capillaries reduced to 2-4 in middle and posterior setigers; hooks ex- VOLUME 92, NUMBER 3 613 Fig. 3. Boccardia pugettensis (paratype, USNM 39567): A, Anterior end in dorsal view, base of left palp only shown; B, Posterior end in ventrolateral view; C, Hooded hook from anterior neuropodium; D, E, Hooded hooks from far posterior neuropodium; F, H, Bristle- topped spines from setiger 5; G, Falcate major spine from setiger 5; I, Notopodium from far posterior segment in dorsal view; J, Notopodial spine and capillary seta from same. 614 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON hibiting wide angle between teeth in anterior and middle setigers (Fig. 3C), this angle reduced in posterior setigers of largest specimens (Fig. 3D), with apical tooth sometimes being entirely lost (Fig. 3E), and hood frequently torn off. Setiger 5 with ventral fascicle of unilimbate capillaries; major spines of 2 types: (1) simple falcate (Fig. 3G); (2) bristle-topped with bristles arising well below expanded tip (Fig. 3F, H). Branchiae on setigers 2, 3, 4, 6 and subsequent segments (Fig. 3A) to end of body, reaching full size by setigers 10-12, where their tips meet middor- sally. Pygidium reduced, much smaller than prepygidial segments, with 4 rounded, glandular lobes, dorsal pair being distinctly smaller than ventral pair (Fig. 3B). Remarks.—The type-specimens of Boccardia pugettensis were previ- ously referred to Polydora (Boccardia) natrix (Soderstrom) by Berkeley and Berkeley (1936; 1952), and, with doubt, by Banse et al. (1968). B. pu- gettensis agrees with B. natrix in having notosetae on setiger 1, an incised prostomium and general similarities in the structure of the major spines of setiger 5. However, a comparison of the holotype of Polydora natrix (SMNH 530) from southern South America, along with new material of the same from off Argentina with the specimens from British Columbia and Washington indicate that 2 distinct species are involved. B. pugettensis has posterior notopodial spines; the hooded hooks in the posterior segments show a reduced angle between the teeth, sometimes with a complete loss of the apical tooth; the expanded ends of the bristle-topped spines of setiger 5 are completely covered by bristles; and the pygidium consists of 4 small glandular lobes. In B. natrix, on the other hand, notopodial spines are lack- ing, the hooded hooks maintain a constant angle between the teeth through- out the body, the expanded ends of the bristle-topped spines of setiger 5 have 2 distinct smooth bosses and the large disclike pygidium is divided into 4 lobes. More detailed comments on the morphology of B. natrix will be included in a study of South American spionids (Blake, in preparation). B. pugettensis appears to be most closely related to B. basilaria Hartman, 1961 from southern California. In B. basilaria, the posterior notopodial spines are heavier than in B. pugettensis, the posterior hooded hooks are falcate and always lose the apical tooth and the pygidium consists of 2 ventral lappets. Moreover, B. basilaria has short anterior branchiae, basally fused to the postsetal lamellae, with those of setigers 2-4 barely extending one-fourth of the distance to the dorsal midline. B. pugettensis, on the other hand, has longer anterior branchiae, not fused basally to the notopodial lamellae, with those of setigers 2—4 extending across the dorsum to near the midline. B. basilaria was incompletely described by Hartman (1961). These comparative comments, based upon an examination of the type-material of VOLUME 92, NUMBER 3 615 B. basilaria (AHF Poly 0637), should serve as an emendation until a more complete description can be prepared. The posterior spines of B. pugettensis, although pointed and subequal in diameter, do not appear to be replacement setae for the long capillaries since they do not end in capillary tips. Distribution.—British Columbia; Washington. In shallow subtidal and in- tertidal sand substrata. Discussion As is evident in the preceding section, there is considerable confusion among different investigators in the interpretation of the notosetae of the posterior setigers in species of the Polydora-complex. In an effort to clarify some of this confusion, a summary of the different forms of posterior no- tosetae found in these species is presented below. 1. Simple fascicles of long and short capillaries, subequal in diameter basally, with or without limbations; the shorter capillaries are developing replacement setae for the longer capillaries. Examples: P. socialis (this paper); P. websteri (Blake, 1971). 2. Long and short capillaries as in (1), with an additional group of short, very thin pointed setae. The latter setae are termed ‘‘needlelike’’ and occur as a group of 15 or more distinct setae as in P. cardalia (this paper) or in a dense flattened packet as in P. flava and P. latispinosa (see Blake and Kudenov, 1978). 3. Long and short capillaries as in (1) with additional short, acicular spines which are not capillary tipped. These spines are usually, but not always, thicker than the accompanying capillaries. Examples: P. caeca (see Fauvel, 1927); P. aciculata (see Blake and Kudenov, 1978). 4. Group of thickened acicular spines, sometimes arranged in a cone or rosette; spines either pointed (awl-shaped) or blunt, with accompanying capillaries usually lacking. Examples: awl-shaped spines—P. caulleryi and P. quadrilobata (see Blake, 1971); long pointed spines—Tripolydora spinosa (see Woodwick, 1964). 5. Capillaries accompanied by 1 or 2 giant, recurved boat hooks, some- times with additional, thickened acicular spines. Examples: P. hoplura (see Fauvel, 1927; Blake and Kudenov, 1978); P. colonia (see Blake, 1971); Boccardiella hamata (see Blake, 1966); B. ligerica (see Blake and Wood- wick, 1971; Light, 1978); B. bihamata (see Blake and Kudenov, 1978). Most of the confusion arises with the first 3 setal arrangements indicated above. The presence of short capillaries among the longer ones has often been interpreted to mean that ‘‘needles’’ or ‘‘short spines’’ are present. Such setae, however, may be merely developing replacements for the longer capillaries. One way to check is to examine several sequential segments. 616 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON If the short setae are lacking on some segments or are highly variable in number relative to the longer capillaries, this suggests a variable replace- ment pattern among the capillaries. True posterior spines or needles will normally be consistent in number and arrangement over several segments. Literature Cited Arai, M. N. 1971. Publications by Edith and Cynl Berkeley.—Jour. Fish. Res. Bd. Canada 28: 1365-1372. Banse, K. 1972. On some species of Phyllodocidae, Syllidae, Nephtyidae, Goniadidae, Apis- tobranchidae, and Spionidae (Polychaeta) from the northeast Pacific Ocean.—Pacific Sci. 26:191—222. Banse, K, K. D. Hobson, and F. H. Nichols. 1968. Annotated list of polychaetes.—Jn U. Lie, A quantitative study of benthic infauna in Puget Sound, Washington, USA, in 1963- 1964. Fiskeridir. Skr. Ser. Havunders. 14:521—548. Berkeley, E. 1927. Polychaetous annelids from the Nanaimo district. 3. Leodicidae to Spion- idae.—Contrib. Canad. Biol. N.S. 3:407—422. Berkeley, E., and C. Berkeley. 1936. Notes on Polychaeta from the coast of western Canada. 1. Spionidae.—Ann. Mag. Nat. Hist., Ser. 10. 18:468—477. , and . 1948. Annelida. Polychaeta Errantia.—Jn Canadian Pacific Fauna. Fish. Res. Bd. Canada 9b(1):100 pp. , and . 1952. Annelida. Polychaeta Sedentaria.—/n Canadian Pacific Fauna. Fish. Res. Bd. Canada 9b(2):139 pp. Blake, J. A. 1966. On Boccardia hamata (Webster), new combination (Polychaeta, Spioni- dae).—Bull. So. Calif. Acad. Sci. 65:176-184. . 1971. Revision of the genus Polydora from the east coast of North America (Poly- chaeta: Spionidae).—Smithsonian Contrib. Zool. 75:32 pp. 1975. Phylum Annelida: Class Polychaeta.—Pp. 151-243 in R. I. Smith and J. A. Carlton (Eds.), Light’s Manual, Intertidal Invertebrates of the Central California Coast. Univ. Calif. Press, Berkeley. Blake, J. A., and J. D. Kudenov. 1978. The Spionidae (Polychaeta) from southeastern Aus- tralia and adjacent areas with a revision of the genera——Mem. Nat. Mus. Victoria 39: 171-280. Blake, J. A., and K. H. Woodwick. 1971. A review of the genus Boccardia Carazzi (Poly- chaeta: Spionidae) with descriptions of two new species.—Bull. So. Calif. Acad. Sci. 70:3 1-42. , and . 1972. New species of Polydora (Polychaeta: Spionidae) from the coast of California.—Bull. So. Calif. Acad. Sci. 70:72-79. Fauvel, P. 1927. Polychetes Sédentaires. Addenda aux Errantes, Archiannélides, Myzosto- maires.—Faune de France 16:494 pp. Foster, N. 1971. Spionidae (Polychaeta) of the Gulf of Mexico and the Caribbean Sea.—Stud. Fauna Curagao Caribb. Islands 36:183 pp. Hartman, O. 1941. Some contributions to the biology and life history of Spionidae from California.—Allan Hancock Pacific Exped. 7:289-324, pls. 45-48. . 1961. Polychaetous annelids from California.—Allan Hancock Pacific Exped. 25:226 pp., 34 pls. ——. 1969. Atlas of the sedentariate polychaetous annelids from California——Allan Han- cock Foundation, Univ. of Southern California, Los Angeles, 812 pp. Light, W. J. 1977. Spionidae (Annelida: Polychaeta) from San Francisco Bay, California: a VOLUME 92, NUMBER 3 617 revised list with nomenclatural changes, new records, and comments on related species from the northeastern Pacific Ocean.—Proc. Biol. Soc. Wash. 90:66—88. . 1978. Spionidae (Polychaeta, Annelida). Invertebrates of the San Francisco Bay Es- tuary System.—The Boxwood Press, Pacific Grove, xii + 211 pp. Mesnil, F. 1896. Etudes de morphologie externe chez les Annélides. I. Les Spionidiens des cotes de la Manche.—Bull. Sci. France Belg. 29:110-—287. Oersted, A. S. 1843. Annulatorum danicorum conspectus, Fasc. 1. Maricolae.—Kgbenhaven, 52 pp., 7 pls: Pettibone, M. H. 1962. New species of polychaete worms (Spionidae: Spiophanes) from the east and west coast of North America.—Proc. Biol. Soc. Wash. 75:77-88. . 1967. Type-specimens of polychaetes described by Edith and Cyril Berkeley (1923- 1964).—Proc. U.S. Nat. Mus. 119(3553):23 pp. Schmarda, L. K. 1861. Neue wirbellose Thiere beobachtet und gesammelt auf einer Reise um die Erde 1853 bis 1857. I. Turbellarien, Rotatorien und Anneliden. Pt. 2.—Leipzig, 164 pp., 22 pls. Sdderstrom, A. 1920. Studien tiber die Polychaetenfamilie Spionidae.—Inaugural-Diss. Upp- sala, 288 pp. Woodwick, K. H. 1963a. Comparison of Boccardia columbiana Berkeley and Boccardia proboscidea Hartman (Annelida, Polychaeta).—Bull. So. Calif. Acad. Sci. 62:132-139. 1963b. Taxonomic revision of two polydorid species (Annelida, Polychaeta, Spio- nidae).—Proc. Biol. Soc. Wash. 76:209-216. . 1964. Polydora and related genera from Eniwetok, Majuro and Bikini Atolls, Marshall Islands.—Pac. Sci. 18:146—159. Tomales Bay Marine Laboratory, Marshall, California 94940. PROC. BIOL. SOC. WASH. 92(3), 1979, pp. 618-633 ADDITIONAL NOTES ON CAVE SHRIMPS (CRUSTACEA: ATYIDAE AND PALAEMONIDAE) FROM THE YUCATAN PENINSULA, MEXICO H. H. Hobbs III Abstract.—The troglobitic shrimps of the Yucatan Peninsula are repre- sented by three species of atyids and a single palaemonid: Typhlatya cam- pechae Hobbs and Hobbs, Typhlatya mitchelli Hobbs and Hobbs, Typhla- tya pearsei Creaser, and Creaseria morleyi (Creaser). These shrimps are found in the states of Campeche, Quintana Roo, and Yucatan and the lo- calities in which specimens have been observed are listed. In addition to a summary of the distribution, observations concerning species variations and life history data are noted along with ecological considerations for each species. Several populations of 7. mitchelli are known to have melanistic representatives and a discussion of the pattern of pigmentation is presented. Introduction.—Creaser (1936, 1938) initiated a study of the troglobitic shrimp fauna of the Yucatan Peninsula when he reported on the crustaceans collected by him during the 1932 University of Michigan Yucatan Expedi- tion. He (1936) described Palaemon morleyi (=Creaseria morleyi) from San Isidro Cave (=Cenote de San Isidro) and Typhlatya pearsei from Balaam Canche Cave [= Grutas de Balankanche]. It was not until 1976 that other shrimp species were reported from the Peninsula. Hobbs and Hobbs (1976), working primarily with specimens collected during 1973 by James R. Reddell (University of Texas at Austin) and associates, described two new species of the genus Typhlatya, T. campecheae and T. mitchelli, and reported many new locality records for the previously known shrimps, T. pearsei (Atyidae) and C. morleyi (Palaemonidae). In late 1974 and mid-1975 Reddell made additional expeditions to the Peninsula and as a result of his efforts, our knowledge of the troglobitic shrimp fauna has increased considerably (see Reddell, 1977). In addition, Holthuis (1977) provided a description and listed new locality records for pigmented specimens of T. mitchelli. This paper treats collections of melanistic specimens of T. mitchelli. Also, synonymies (only additions to those by Hobbs and Hobbs, 1976) are pre- sented for the four troglobitic shrimps occurring in the State of Yucatan. Summaries of the distribution and observations concerning variations are noted along with ecological considerations for each of them. Life history data are tabulated in Table 2 and reflect the bias in sampling since 1932. VOLUME 92, NUMBER 3 619 The reader is referred to Hobbs and Hobbs (1976) for descriptions of the atyids and to Holthuis (1952) for a description of the palaemonid, C. mor- leyi. For information concerning geology, physiography, and cave and karst development in the Yucatan Peninsula, see Butterlin and Bonet (1936), Finch (1973), West (1964), Isphording (1975), and Isphording and Wilson (1973). Acknowledgments.—I am grateful to James R. Reddell for not only mak- ing specimens available for examination, but also for his assistance in plot- ting cave entrance locations and for information concerning the ecology of the shrimps. I am also indebted to Lipke B. Holthuis of the Rijksmuseum van Na- tuurlijke Historie, Leiden, Netherlands, for informing me of his recent work on the cave shrimps from Mexico. I extend my appreciation to Horst Wilk- ens of the Zoological Institute of the University of Hamburg, West Ger- many, for notifying me of the disposition of the shrimp specimens he col- lected during the third Biospeleological Expedition to Mexico by the Accademia Nazionale dei Lincei. I am most grateful to my father, Horton H. Hobbs, Jr., for initially di- recting me in the study of these shrimps and for his criticisms and aid in the preparation of this manuscript. Listed below, alphabetically, are the collectors of the specimens exam- ined (initials are used to identify them in the species treatments). Especial thanks are extended to those persons who spent considerable time in the Peninsula caves collecting shrimps. Francis E. Abernethy—F.E.A., E. W. Andrews—E.W.A., R. Argano— R.A., Mary Butterwich—M.B., Jerry W. Cooke—J.W.C., E. P. Creaser— E.P.C., Deborah Denson—D.D., Linda Elliott—L.E., M. Cardenas Fi- gueroa—M.C.F., Andy Grubbs—A.G., F.G. Hall—F.G.H., Masaharu Ka- wakatsu—M.K., David McKenzie—D.M., Martha H. McKenzie—M.H.M.., Robert W. Mitchell—R.W.M., S. R. Mitchell—S.R.M., Sharon A. Mitch- ell—S.A.M., Stuart Murphy—S.M., A. S. Pearse—A.S.P., Terry Raines— T.R., James R. Reddell—J.R.R., William H. Russell—W.H.R., E. H. Sal- lee—E.H.S., Reynaldo Solis—R.S., B. F. Osorio Tafall—B.F.O.T., Su- zanne Wiley—S.W., and H. Wilkens—H.W. Suborder Natantia Section Caridea Family Atyidae Genus Typhlatya Typhlatya campecheae Hobbs and Hobbs Typhlatya campecheae Hobbs and Hobbs, 1976:6, 7 (fig. 3), 8, 9 (fig 4), 10, 14-16, 20 (Appendix), 21 (Map 1).—Buden and Felder, 1977:114.— 620 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Hobbs, Hobbs, and Daniel, 1977:i11, 6, 8, 11, 12, 24, 32, 33 (fig. 7), 34 (fig. 8).—Reddell, 1977:230, 240 (Table 1) 241, 243, 249.—Holthuis, 1977:174. Type-locality.—Grutas de Xtacumbilxunam (=Grutas de Ixtacumbilxu- nan; Grutas de Xtacumbilxunan), 2 km SW Bolonchenticul, Campeche, Mexico. Distribution. —Campeche, Mexico. Known from the type-locality and from Cenote de Cantemo, | km N Cantemo, Campeche, Mexico. Specimens examined.—217 specimens have been examined from Cam- peche, Mexico; locality numbers in parentheses correspond to the locality numbers in Table 1 and Figure 1. (5) Type-locality, 1¢, 42, 22 ovig., 19 April 1973, and 40d, 1092 and 21 juv., 13 May 1973, M.B., D.M., M.H.M.., S.M., and J.R.R., coll. (2) Cenote de Cantemo, 18°, 18 Dec. 1974, L.E., D.M., and J.R.R., coll.; 222, 31 July 1975, A.G., D.M., J.R.R., and S.W., coll. Ecology.—T. campecheae has been found only in lentic environments in these two caves. In the type-locality it frequented guano-floored pools, and in Cenote de Cantemo it inhabited a pool 1.5 m deep and 3 m in diameter, floored with organic debris, the surface being covered with a thin layer of oil (see Reddell, 1977 for descriptions of the caves and for faunal associates; see Table 2 for life history data). Color.—Preserved specimens are translucent to white, totally lacking pig- ments. Variations.—Hobbs and Hobbs (1976:10) discussed variations in the pop- ulation from the type-locality. No additional variations were observed in the specimens examined from Cenote de Cantemo. Typhlatya mitchelli Hobbs and Hobbs Typhlatya mitchelli Hobbs and Hobbs, 1976:2, 3 (fig. 1), 4, 5 (fig. 2), 6, 10, 14, 15, 16, 20 (Appendix), 21 (Map 1).—Buden and Felder, 1977:114.— Hobbs, Hobbs, and Daniel, 1977:iii, 6, 8, 11, 12, 25, 39, 40 (fig. 12), 41 (fig. 13), 45.—Reddell, 1977:230, 239, 240 (Table 1), 251, 255, 256, 260, 262, 263, 268, 279, 281, 289, 291, 292, 293.—Holthuis, 1977: 173-176 (fig. la-i), 177, 178 (fig. 2), 179-181, 194. Type-locality.—Cenote Kabahchén (=Actun Caba Chen), in the town of Muni, Yucatan, México. Distribution.—This species is known from the states of Quintana Roo and Yucatan (Northwestern and Northeastern Coastal Plain physiographic dis- tricts). Specimens examined.—A total of 55 specimens, the majority females, have been collected from the states of Quintana Roo and Yucatan; those specimens marked by an asterisk were not examined by the author; locality VOLUME 92, NUMBER 3 621 aim 42 @30 aoe 29 Xo7 a 55 // 28 y VX fi *. 25 y21 226 4lS 54 43 * YAXCABA @r- a Tm BTp @cm ¥™ms Tp OtTméscm % Tp &Cm 9 @ Tm &Tp& Gm BB Shrimp sighted oe EOsON Ln FELIPE CARRILLO Fig. 1. Distribution of the troglobitic shrimps (Atyidae and Palaemonidae) in the Yucatan Peninsula, Mexico. Location of the entrance to Cueva Coop (No. 13) not shown. Tc = Typhlatya campecheae; Tm = Typhlatya mitchelli; Tp = Typhlatya pearsei; and Cm = Creas- eria morleyi. numbers in parentheses correspond to the locality numbers in Table 1 and on Figure 1. QUINTANA ROO: (6) Curva del Fermin, 3 km E Pamul at Rancho San Martin, 22, 3 July 1975, A.G., D.M., and J.R.R., coll. (7) Actun Ha, located in the ruins of Coba, 22, 30 June 1975, A.G., D.M.., J.R.R., and S.W., coll. YUCATAN: (14) Cenote de la Culebra, 20 m W of México Hwy 180, midway between km 175 and 176, 1°, 1J, 27 March 1973, J.R.R., coll. (15) Cenote Chun Kopac (=Cenote Acanceh) Acanceh, 1°, 8 October 1974, J.R.R., coll. (20) Actun Kaua, | km S Kaua, 1°, 11 December 1974, A.G., D.M., J.R.R., and S.W., coll. (23) Type-locality, 52 plus ho- lotype, 1 August 1973, M.K., R.W.M., and J.R.R., coll. (25) Cenote Cal- chuhuim (=? Cenote Calchuntunil), 2 km E Hacienda San Bernardo, 6°, 15 October 1974, J.R.R., R.S., and S.W., coll. (32) Cenote del Pochote (=Cueva de Caxcuy, Cueva del Ponte, Cueva del Pochote), 10 km NW Muna, 82°, 4 November 1974, D.M., J.R.R., and S.W., coll. (38) Cenote Chen Mul (=Cenote Ch’en Mul), located at the base of the great pyramid in the Ruins of Mayapan, 22, 26 April 1973, M.B., D.M., M.H.M., and 622 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON J.R.R., coll. (40) Grutas de Tzab-Nah (=Grutas de Dzab-Nah), 2 km S Tecoh, 32, 6 August 1973, D.D., R.W.M., S.A.M., and S.R.M., coll. (44) Cenote Aka Chen, | km E Tixcancal, 32, 2 April 1973, M.B., D.M., M.H.M., S.M., and J.R.R., coll. (45) Cenote de Orizaba (=Cueva de Ori- zaba), 8 km S Buenaventura, 12, 1 April 1973, M.B., D.M., M.H.M., S.M., J.R.R., coll. (47) Cenote Sodzil, 5 km W Sucopo, 52, 31 March 1973, D.M.., M.H.M., S.M., and J.R.R., coll. (50) Cenote Zaci*, near the highway from Tizimin to Colonia Yucatan and El Cuyo, 36 km from Tizimin, 3 speci- mens*, March 1975, H.W., coll. (51) Cenote de la Paca, 7 km E Tikuch, 22, 11 April 1973, S.M., coll. (52) Cenote Xkeken, 3 km N Dzit-Nut, 4°, 10 December 1974, D.M. and J.R.R., coll; 1 specimen*, March 1975, H.W., coll. (53) Cenote Xtacabiha (=Cenote Xalau = Cenote de Xtacabiha) 9 km NNE Tikuch, 22, 11 April 1973, M.H.M., S.M., and J.R.R., coll.; 12, 6 July 1975, A.G. and J.R.R., coll. Ecology.—Typhlatya mitchelli has been collected from 2 localities in Quintana Roo and 15 from Yucatan; ‘“‘atyid shrimp,’ among which this species may have been represented, have been observed but not captured from four additional cenotes in Yucatan (see Reddell, 1977:257, 262, 279, 290). In all localities shrimps were living in lentic situations varying from very small shallow (less than 0.3 m depth) pools to extensive underground lakes (greater than 2.0 m depth). Pearse (1938:12) reported an ‘‘inner pool’’ of Actun Kaua to have a temperature of 24.5°C on 17 June 1936. Generally the pools were in total darkness, but 7. mitchelli occurs in entrance areas that receive direct light from the surface (e.g. Grutas de Tzab-Nah). Sub- Strates of the pools consisted of guano, silt, organic debris, rocks and com- binations thereof. Shrimps were found on the substrate, ‘“‘hanging’’ from submerged walls along the edges of lakes, free-swimming, and among dense, branched root systems hanging down from the roofs of cenotes. Holthuis (1977:181) states, ‘‘The finely branched root systems of trees or other plants exposed in the water are among the favourite [sic] habitats of small atyids all over the world, whether subterranean or epigean.”’ For a discussion of the fauna found in association with T. mitchelli see Reddell (1977) and Table 1. See Table 2 for life history data. Color.—Hobbs and Hobbs (1976:4) in describing T. mitchelli noted that all the preserved specimens they examined (25) were ‘‘translucent to white.’’ On 6 July 1975 James Reddell et al. made an additional collection of shrimps from Cenote Xtacabiha. Unlike those captured on their first visit (11 April 1973), these specimens were pigmented. Shortly after I examined the most recent collection and also a pigmented series from Cenote Xkeken, I received a letter from Horst Wilkens in which he mentioned that he had collected pigmented specimens of T. mitchelli from two localities in Yucatan and had sent them to Lipke Holthuis. Correspondence with Dr. Holthuis indicated that he was treating these specimens in his manuscript on the VOLUME 92, NUMBER 3 623 Fig. 2. Lateral view of melanistic female specimen of Typhlatya mitchelli from Cenote Xtacahiba showing color pattern. subterranean shrimps of the Italian Mexico Expedition 1973 (see Holthuis, 1977:175—181). In this paper, Dr. Holthuis not only indicated the coloration of this shrimp but also figured and described the male of the species. Figure 2 is a lateral view of a melanistic female specimen (4.2 mm cara- pace length) from Cenote Xtacabiha, showing the characteristic “‘dark chro- matophore-like specks’’ (Holthuis, 1977:180) characteristic of specimens from the three cenote localities. The pigment appears greyish to black in preserved specimens (brown in a few specimens) and is particularly distinct along the ventral margin of the carapace, the rostrum, the eye-remnant (pigmented but not faceted), the first abdominal tergum, and the antennular peduncle (see Holthuis, 1977:180 for additional remarks concerning pig- mentation). It is noteworthy that after being preserved in 70% ethanol for more than five years, the pigment is still easily observed in the specimens. Variations.—No significant variations were noted in the specimens ex- amined from localities other than those listed in Hobbs and Hobbs (1976). 624 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Table 1. Distribution of troglobitic shrimps from the Yucatan Peninsula, Mexico. Palaemo- Atyidae rida WM, T. C. Localities campecheae_ mitchelli T. pearsei morleyi CAMPECHE . Cenote de Bolchen — — = an . Cenote de Cantemo + — = se . Grutas de Monte Bravo — = + . Grutas de San Antonio — = ie . Grutas de Xtacumbilcunam + = a: £3 QUINTANA ROO 6. Cueva del Fermin ~ + 4 Jt 7. Actun Ha = + s p! 8. Pozo de San Martin ~ — + a" 9. Cenote de Juan Coh - _ 4. tt 10. Cenote de Las Ruinas — — = ae + + + nA bh WN = 11. Cenote de Santo Domingo — _ of 12. Cenote de Tos Virlol — = ie 13. Cueva Coop = = ae YUCATAN 14. Cenote de la Culebra — + = ae 15. Cenote Chun Kapoc = ot a 2 16. Cenote de San Diego = ? ? 17. Cenote de Hoctun = = ae ar 18. Cenote X-ebiz = = = a 19. Cueva Chac Mol = = == ar 20. Actun Kaua eT ar 21. Cenote de las Abejas = 22. Cenote de Sambula = 23. Cenote Kabahchen = 24. Cenote Calchum = = 25. Cenote Calchuhuim — + = 26. Cueva Amil = = == 27. Cenote de San Isidro ~ = st 28. Cenote de San Jose — -- = 29. Cenote de Huolpoch = = = 30. Cenote de Sambula = = = 31. Cenote de Kankirixche = = = 32. Cenote del Pochote — + 3 33. Actun Xpukil = = = 34. Actun Gongora — = = 35. Cenote Nohchen ~ _ + 36. Actun Chac - — + 37. Actun Okobichen — — + 38. Cenote Chen Mul = 39. Cenote de Telchaquillo = 40. Grutas de Tzab-Nah — 41. Cueva de Santa Elena = = stig “gee i? Moria a he ese ar aa) Oe eel ar? Gao he VOLUME 92, NUMBER 3 625 Table 1. Continued. Palaemo- Atyidae Prdae ’ Ike C. Localities campecheae_ mitchelli T. pearsei morleyi 42. Pozo de Santa Elena - 43. Grutas de Balankanche — 44. Cenote Aka Chen — 45. Cenote de Orizaba = 46. Cenote de los Pinos = 47. Cenote Sodzil — 48. Cueva Sodzil — 49. Cueva Xconsacab - 50. Cenote Zaci — 51. Cenote de la Paca _ 52. Cenote Xkeken ~ 53. Cenote Xtacabiha — 54. Cenote Yunchen — 55. Cueva Luchil — - - + + l+ot+4 1 | | +++ 41 Holthuis (1977) described specimens from Cenote Zaci and Cenote Xkeken and indicated that the endopod of the first pleopod of the male is “‘oval with some hairs in the outer half.’’ He (p. 180) described the appendix masculina of the male second pleopod as ‘‘short and oval, but still longer than the appendix interna; it ends in three rather strong bristles.”’ Typhlatya pearsei Creaser Typhlatya pearsei.—Holthuis, 1977:173-176, 180, 181, 194, fig. 1j.—Red- delle 1977-230. 239=241 243. 247, 251-252, 258,260, 262, 263. 266, 277, 278, 281, 283, 285, 293, Table 1.—Buden and Felder, 1977:112.—Hobbs and Hobbs, 1976:1, 10-16, 20, 21, fig. 5, map 1, Appendix.—Hobbs, Hobbs, and Daniel, 1977:1ii, 6, 7, 9, 12, 15, 24, 39, 43-45, figs. 3, 15, and 16. Type-locality.—Grutas de Balankanche (=Cueva de Balaam Canche), 4 km E Chichén Itza, Yucatan, México. Distribution.—T. pearsei is the most widely distributed atyid on the Yu- catan Peninsula, being known from 21 localities in the Northeastern and Northwestern Coastal Plain, the Sierra de Ticul, and the Sierra de Bolon- chén physiographic districts in the states of Campeche, Quintana Roo, and Yucatan. Specimens examined.—222 specimens have been collected from 23 lo- calities in Yucatan, México; locality numbers in parentheses correspond to the locality numbers in Table 1 and Figure 1; specimens marked with an 626 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON asterisk were not examined by the author. CAMPECHE: (3) Grutas de Monte Bravo, 10 km NW Cantemo, 3°, 19 December 1974, L.E., D.M., and J.R.R., coll. (4) Grutas de San Antonio, 10 km NNE Bolonchenticul, 26,72, 3 November 1974, D.M., J.R.R., and S.W., coll.; 152, 23-24 No- vember 1974, D.M., J.R.R., and S.W., coll. QUINTANA ROO: (6) Cueva del Fermin, 42, 3 July 1975, A.G., D.M., and J.R.R., coll. (8) Pozo de San Martin, 2.5 km E Pamul at Rancho San Martin, 42, 2 July 1975, J.R.R., coll. (9) Cenote de Juan Coh, Felipe Carrillo Puerto, 92, 1 juv., 4 July 1975, A.G., D.M., and J.R.R., coll. (11) Cenote de Santo Domingo, 5 km ENE kilometer 50, 82, 28 July 1975, A.G., D.M., and J.R.R., coll. (12) Cenote de Tos Virlol, 13 km S Senor, 12, 4 July 1975, A.G. and J.R.R., coll. (13) Cueva Coop, near village of Pamul, 120 km NE Felipe Carrillo Puerto, 1 specimen*, March 1975, H.W., coll. YUCATAN: (17) Cenote de Hoctun (=Cueva de Hoctun), 1 km W Hoctun, 72, 16 March 1973, M.B., D.M., M.H.M., S.M., J.R.R., coll.; 1 specimen*, March 1975, H.W., coll. (20) Actun Kaua, | km S Kaua, 32, 20-21 October 1974, D.M., J.R.R., and S.W., coll. (21) Cenote de las Abejas, 2 km N Kopoma, 1d, 22, 16 April 1973, D.M. and S.M., coll. (23) Cenote Kabahchén (=Actun Caba Chén), Mani, 52, | August 1973, D.M., S.M., and J.R.R., coll. (24) Cenote Cal- chun, | km E Hacienda San Bernardo, 1d, 16 April 1973, D.M., S.M., and J.R.R., coll. (27) Cenote de San Isidro (=Cueva de San Isidro, Cenote de El Retiro), southern end of Calle 66, Merida, 1°, 21 March 1973, S.M. and J.R.R., coll. (32) Cenote del Pochote, no specimens examined, reported from this locality by Cardenas (1950:156) and Nicholas (1962: 173). (35) Cen- ote Nohchén, Sacalum, 4¢, 292, 18 June 1975, A.G., D.M., and J.R.R., coll. (36) Actun Chac (=Gruta de Chac), S of Kabah, 12°, 24 November 1962, E.W.A., coll. (37) Actun Okobichén, 8 km SW Santa Elena, 1d, 9°, 15 November 1974, D.M., J.R.R., and S.W., coll. (40) Grutas de Tzab-Nah, 42,4 juv., 22 April 1973, D.M. and J.R.R., coll. (41) Cueva de Santa Elena (=Cueva or Cenote de los Camarones), 4.8 km S Telchac Puerto, 1°, 31 July 1932, E.P.C., coll.; 3d, 242, 6 juv., 22 March 1973, J.R.R., coll. (43) Type-locality, 62, 24 June 1932, E.P.C. and A.S.P., coll.; 122, 2 juv., 28 June 1932, E.P.C. and A.S.P., coll.; 302, 1 juv., 30 June 1932, E.P.C., coll. (53) Cenote Xtacabiha, 8°, 11 April 1973, D.M., M.H.M., S.M., and J.R.R., coll. (55) Cueva Luchil (=Cenote Luchil) 8 km SSE Mérida, 12, 25 July 1932, E.P.C., coll. (Creaser’s label for the specimens reads ‘‘Tuchil-cave on Tixcacal Hacienda, 14 km SE Merida, Yucatan’’ and is believed to be the same location as Cueva Luchil.) Reddell (1977) mentions three caves (see Table 1) in which atyid shrimps were observed. It is highly improbable that they were members of T. cam- pecheae, but it is likely that they belonged to either 7. mitchelli or T. pearseli. Ecology.—Typhlatya pearsei has been collected from 2 localities in Cam- VOLUME 92, NUMBER 3 627 peche, 6 caves in Quintana Roo, and 15 localities in Yucatan. This species is found in lentic habitats ranging from small pools (less than 0.5 m in depth) to deep lakes (greater than 8 m). These bodies of water are generally floored with silt or bat guano, and in Cenote de San Isidro the water is contaminated with human trash and feces. Reddell (1977:258) indicated that Cueva Luchil was ‘“‘badly polluted by oil and debris. Piles of human feces are abundant on the rocks below the entrance.’’ Water temperatures have been reported by Hall (1936), Pearse (1938), Robles Ramos (1950), and by Reddell (1977) for various caves, ranging from 23.5°C to 28.3°C. Hall (1936:12) presented chemical and physical data for the type-locality of this shrimp and for Cueva de Santa Elena; pH values were 7.4 and 6.8, respectively and those of dissolved oxygen, 6.51 and 0.78 mg 171, respectively. These data are similar to those he obtained from other localities in the region (see Hall, 1936:11). Hence, it can be concluded that 7. pearsei, and other aquatic cavernicoles from the Peninsula, are able to tolerate oxygen levels that are well below saturation (i.e. 10%-—80% levels). Typhlatya pearsei is generally found in total darkness but occasionally individuals are seen in entrance areas where pools receive either direct or indirect light from the surface. See Reddell (1977) for other faunal elements and for a detailed description of the caves. See Table 1 for shrimp associates and Table 2 for life history data. Color.—Preserved specimens are translucent to white; Holthuis (1977:175) noted that when alive they “‘are of a light, whitish, colour [sic], being somewhat opaque, not transparent.” Variations.—Generally, specimens examined from localities other than those reported by Hobbs and Hobbs showed no significant morphological variations from published descriptions of the species. However, several males from Cenote Nohchén in Yucatan possessed an atypical second pleo- pod (Fig. 3A, D). The appendix masculina is much longer than the appendix interna, as the latter is proportionately shorter than that exhibited by more ‘‘typical’’ males from other localities. In addition, only 5 apical and sub- apical spines are present on the apex of the appendix masculina as compared to approximately 16 in specimens from other caves. Family Palaemonidae Genus Creaseria Creaseria morleyi (Creaser) Palaemon morleyi.—Hobbs and Hobbs, 1976:1.—Hobbs, Hobbs, and Dan- tel, 197 727, 9: Creaseria morleyi.—Holthuis, 1977:173, 186, 187, fig. 5a—c.—Reddell, 1977:230, 239, 240, 241, 243, 247, 251-253, 255, 258, 262-264, 266-268, 273, 274, 277, 281, 283, 285, 291-293, Table 1.—Hobbs and Hobbs, 628 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 1976:1, 16-21, figs. 7, 8, Appendix, Map 2.—Hobbs, Hobbs, and Daniel, 1977:i11, 6, 9, 12, 25, 46-49, figs. 18, 19. Type-locality.—Cenote de San Isidro (Cueva de San Isidro, Cenote de El Retiro), near southern end of Calle 66 in Mérida, Yucatan, México. Distribution.—Even though its range does not greatly exceed that of Typhlatya pearsei, C. morleyi is probably the most common cavernicolous shrimp within the Peninsula, being known from the states of Campeche, Quintana Roo and Yucatan (the Northwestern and Northeastern Coastal Plains, the Sierra de Ticul, and the Sierra de Bolonchén physiographic dis- tricts). Specimens examined.—101 specimens were taken from 32 localities in the states of Campeche, Quintana Roo, and Yucatan; locality numbers in parentheses correspond to the locality numbers in Table 1 and Figure 1; specimens marked with an asterisk were not examined by the author. CAM- PECHE: (1) Cenote de Bolchén, 3 km S San Antonio Sacabchén, 36, 6°, 30 October 1974, D.M., J.R.R., and S.W., coll. (3) Grutas de Monte Bravo, 22, 19 December 1974, L.E., D.M., and J.R.R., coll. (4) Grutas de San Antonio, 16, 3 November 1974, D.M., J.R.R., and S.W., coll. QUINTANA ROO: (6) Cueva del Fermin, 1¢, 12, 3 July 1975, A.G., D.M., and J.R.R., coll. (8) Pozo de San Martin, 16, 52, 2 July 1975, J.R.R., coll. (9) Cenote de Juan Coh, 446, 62, 4 July 1975, A.G., D.M., and J.R.R., coll. (10) Cenote de las Ruinas, 6 km ENE Polyuc, 16,62, 29 July 1975, A.G., D.M., J.R.R., and S.W., coll. (11) Cenote de Santo Domingo, 2d, 42, 29 July 1975, A.G., D.M., and J.R.R., coll. (12) Cenote de Tos Virlol, 3d, 82, 4 July 1975, A.G. and J.R.R., coll. (13) Cueva Coop, | specimen*, March 1975, H.W.., coll. YUCATAN: (14) Cenote de la Culebra, 22, 3 August 1973, F.E.A. and R.W.M., coll. (17) Cenote de Hocttin, 22, March 1969, T.R., coll.; 2 specimens*, spring 1970, H.W., coll.; 32, 16 March 1973, M.B., D.M., M.H.M., S.M., and J.R.R., coll.; 12, 29 April 1973, R.W.M. and J.R.R., coll.; 22, 8 August 1973, F.E.A., D.D., M.K., R.W.M., S.A.M., S.R.M., coll. (18) Cenote X-ebiz, near Hoctun, 1°, 26 April 1971, E.H.S., coll. (19) Cueva Chac Mol*, near Tohil, reported by Creaser (1938:163). (21) Cenote de las Abejas, 1d, 16 April 1973, D.M. and S.M., coll. (23) Cenote Ka- bahchén, 1d, 12, 1 August 1973, M.K., R.W.M., and J.R.R., coll. (25) Cenote Calchuhuim, 1¢, 12, 15 October 1974, J.R.R., R.S., and S.W., coll. (26) Cueva Amil (=Cenote Amil)*, on Hacienda Tixcacal, 2 km E Merida, reported by Creaser (1936:128). (27) type-locality, 42 , 21 March 1973, S.M. and J.R.R., coll. (30) Cenote de Sambula (=Cueva de San Bulha)*, Motul, reported by Creaser (1936:128). (31) Cenote de Kankirixché*, on Rancho San Isidro Kankirixché, 10 km NW Muna, reported by Reddell (1977:268). (32) Cenote del Pochote, 23, 4 November 1974, D.M., J.R.R., and S.W., coll.; 12*, March 1975, H.W., coll. (33) Actin Xpukil (=Actun Spukil, VOLUME 92, NUMBER 3 629 Table 2. Life history data for troglobitic shrimps of the Yucatan Peninsula (month, sex, size, range and number of specimens collected since 1932). T. campecheae T. mitchelli T. pearsei T. morleyi January — — —- 1 specimen*** February — — — —_ March —_ 6° aig 13 1 Juv. SPR Se Ise 4 specimens** 6 Juv. 1 specimen** 2 specimens** April NG) 10 2 26 36 4.2 9° 14 9 a 4 Juv. May 40 3 — — 2 specimens** 109 ° 21 Juv June — De 46 specimens*** 12 3 Juv. specimens** July DDS 32 28 ¢ 126 1 Juv. 30 2 specimens*** specimens*** August — oy 529 13 y 2 4 specimens**** September — — — — October — 12 352 4¢6 2 November — 8° 34 3G 329 ie 3 Juv. December 18 @ Sys ao ge) Total number 2A 515) 222 101 of specimens Carapace length 3d —3.0-3.9 = d—3.6-3.8 d—14.0-20.3 (range) a Sela 2—3.4-4.8 ?—3.1-5.2 oe 123-2910 Ue vee Juyv— 32.3 Juv.—S5.6—9.0 * Ovigerous (approximately 0.8 x 11.1 mm—egg size). ** See Holthuis, 1977. *** See Creaser, 1936. a SCeILeatse-alo56- 630 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Grutas de Calcehtok, Cueva de Oxkintok)*, in Sierra de Ticul, 3 km S Calcehtok, reported by Creaser (1938:163). (34) Actun Gongora (=Actun Gongurrah, Cueva de Gorgosa)*, 1.5 km S Oxkutzcab, reported by Creaser (1938:163). (36) Actin Chac, 2 juv., 24 November 1962, E.W.A., coll.; 19, 15 December 1962, E.W.A., coll.; 12, 1 juv., 16 November 1974, D.M.., J.R.R., and S.W., coll. (40) Grutas de Tzab-Nah, 1d, 22, 22 April 1973, D.M. and J.R.R., coll. (42) Pozo de Santa Elena, 4.8 km S Telchac Puerto, 16,22, 22 March 1973, S.M. and J.R.R., coll. Hobbs and Hobbs (1976:20) erroneously reported this species from Cueva de Santa Elena and listed the locality of Pozo de Santa Elena as **5 km S Telchac Puerto”’ (p. 16) which, in fact, is the location for Cueva de Santa Elena. (43) Grutas de Balan- kanche, specimens* from Pearse (1936:128); 4 specimens*, 23 August 1973, R.A., coll.; 1¢, “‘Ist half, 1961,°> E.W.A., coll. (48) Cueva Sodzil, 5 km W Sucopo and 50 m from Cenote Sodzil, 3°, 31 March 1973, D.M., M.H.M.., S.M., and J.R.R., coll. (49) Cueva Xconsacob, near Tizimin, 11 August 1936, A.S.P., sight record by Creaser (1938:163). (51) Cenote de la Paca, 1d, 12, 11 April 1973, S.M., coll. (54) Cenote Yunchén (=Cueva Yunchén)*, Libre Union, 11 July 1936, A.S.P., reported by Creaser (1938:163). Reddell (1977) mentioned five localities in which palaemonids were observed but not captured (see Table 1). Ecology.—Creaseria morleyi occurs in quiet water habitats ranging from small, shallow pools (less than 0.5 m in depth) and natural wells (5 m deep) to large, deep lakes (greater than 3 m in depth). These bodies of water are floored by guano deposits, rich organic silt, debris, and rocks. Hall (1936), Hubbs (1936), Pearse (1938), Robles Ramos (1950), and Reddell (1977) re- ported water temperature in 11 localities and these values ranged from 23.5°— 28.3°C. Shrimp were found in total darkness as well as in entrance areas receiving direct and indirect light. Hall (1936) presented physical and chem- ical data for some of the caves inhabited by C. morleyi: pH ranged between 6.8 and 7.4 and oxygen concentration, from 0.78 to 6.51 mg 1~!. These shrimp have been observed to crawl over the bottom of the substrate as well as to ‘‘swim’’ throughout the water column (Creaser, 1936 and Reddell, 1977). Creaser also reported that C. morleyi is quite sensitive to water vibration and that the species is cannibalistic. Holthuis (1977:187) states that “‘these animals are very agressive [sic], if two or more specimens are placed together in a smallish container they will attack and mutilate each other.’’ See Table 2 for life history data. — Fig. 3. A. Second pleopod of male Typhlatya pearsei from Cenote Nohchen; B. Lateral view of anterior region of male Creaseria morleyi from Cenote Calchuntunil; C. Lateral view of anterior region of male Creaseria morleyi from Grutas de San Antonio; D. Appendices masculina and interna of male Typhlatya pearsei from Cenote Nohchén. VOLUME 92, NUMBER 3 D 0.1 B 5.0 C 5.0 A 0.1 Scales” in mm 632 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Color.—Preserved specimens are translucent to white. Variations.—This appears to be a rather variable species: particularly conspicuous are the differences noted in the second pleopod of the male and the rostrum of both sexes. The appendix interna may or may not extend to the apex of the appendix masculina, and the number of spines on the latter varies from 19 to 27. The rostrum is extremely variable, ranging from long, thin, nearly straight and not expanded dorsoventrally (Fig. 3B) to short, thick, and vaulted (Fig. 3C). Rostral spination varies in number (from 6-10 dorsally and 0-3 ventrally) and relative position on dorsal and ventral sides, and the apex of the rostrum may be acutely spined or somewhat rounded. Generally, if the apex of the rostrum reaches or extends beyond the antennular peduncle, then the spines are acute; otherwise they tend to be rounded, little more than ‘“‘bumps.’’ As Holthuis (1977:187) indicates, ‘‘the eye is not always perfectly bullet-shaped as described, but often shows a distinct tubercle on the outer surface somewhat behind the apex . . .”’ and in ‘‘larger specimens the crenulated tooth of the fixed finger of the second leg . . . is distinct, the cutting edge of the dactylus bears a triangular tooth distally of the tooth of the fixed finger.’’ Literature Cited Buden, Donald W., and Darryl L. Felder. 1977. Cave shrimps in the Caicos Islands.—Proc. Biol. Soc. Washington 90: 108-115. Butterlin, J., and F. Bonet. 1963. Mapas geologicos de la Peninsula de Yucatan. I-Las for- maciones cenozoicas de la parte mexicana de la Peninsula de Yucatan.—Ingin. Hidra. México 17:63-71. Creaser, E. P. 1936. Crustaceans from Yucatan.—Carnegie Inst. Washington Publ. 457:117- NS), . 1938. Larger cave crustacea of the Yucatan Peninsula.—Carnegie Inst. Washington Publ. 491:159-164. Finch, W. A., Jr. 1973. The karst landscape of Yucatan.—Ph.D. Dissertation Univ. Illinois, Univ. Microfilms, Inc., Ann Arbor, Michigan, 168 pp. Hall, F. G. 1936. Physical and chemical survey of cenotes of Yucatan.—Carnegie Inst. Wash- ington Publ. 457:5-16. Hobbs, Horton H., Jr., H. H. Hobbs III, and Margaret A. Daniel. 1977. A review of the troglobitic decapod crustaceans of the Americas.—Smithson. Contrib. Zool. 244: 1-183. Hobbs, H. H. III, and Horton H. Hobbs, Jr. 1976. On the troglobitic shrimps of the Yucatan Peninsula, Mexico (Decapoda: Atyidae and Palaemonidae).—Smithson. Contrib. Zool. 240: 1-23. Holthuis, L. B. 1952. A general revision of the Palaemonidae (Crustacea Decapoda Natantia) of the Americas, II: The Subfamily Palaemoninae.—Allen Hancock Found. Occ. Pap. 12:1-396. ———. 1977. Cave shrimps (Crustacea Decapoda, Natantia) from Mexico. Part IIJ].—Jn Sub- terranean Fauna of Mexico. Accad. Nazionale Lincei 171:173-195. Hubbs, C. L. 1936. Fishes of the Yucatan Peninsula.—Carnegie Inst. Washington Publ. 457:157—287. Isphording, W. C. 1975. The physical geology of Yucatan.—Trans. Gulf Coast Assoc. Geol. Soc. 25:231-262. VOLUME 92, NUMBER 3 633 , and E. M. Wilson. 1973. Weathering processes and physical subdivisions of northern Yucatan.—Proc. Assoc. Amer. Geograph. 5:117-121. Pearse, A. S. 1936. Results of survey of the cenotes in Yucatan.—Carnegie Inst. Washington Publ. 457:17-28. . 1938. Fauna of the caves of Yucatan, Introduction.—Carnegie Inst. Washington Publ. 491:1-17. Reddell, James A. 1977. A preliminary survey of the caves of the Yucatan Peninsula.—Assoc. Mexican Cave Stud. Bull. 6:215—296. Robles Ramos, R. 1950. Los recursos naturales de Yucatan: I.—Apuntes sobre la morfologia de Yucatan.—Bol. Soc. Mexicana Geogr. Estadist. 69:27—106. West, R. C. 1964. The natural regions of Middle America.—Pp. 363-383 in R. C. West, ed., Handbook of Middle American Indians, vol. 1: Natural environment and early cultures. Univ. Texas Press, Austin, 363—383 pp. Department of Biology, Wittenberg University, Springfield, Ohio 45501. PROC. BIOL. SOC. WASH. 92(3), 1979, pp. 634-639 THREE NEW STOMATOPOD CRUSTACEANS FROM THE PACIFIC COAST OF COSTA RICA Raymond B. Manning and Marjorie L. Reaka Abstract.—Gonodactylus albicinctus, G. costaricensis, and Nannosquilla canica are described from specimens collected off the Pacific coast of Costa Rica. Among the stomatopod Crustacea collected by one of us (M. L. R.) while participating in the R/V Searcher Expedition to Costa Rica in 1972 are three species, two in Gonodactylus and one in Nannosquilla, which appear to be undescribed. These three species are characterized below, so that names can be used in a review of the distributional ecology of Costa Rican sto- matopods which is in preparation. All of these specimens have been deposited in the Division of Crustacea, National Museum of Natural History, Smithsonian Institution. For each Species specimens other than the holotype are paratypes. Measurements are in millimeters; total length is abbreviated to TL. Original references for other species discussed below can be found in Schmitt (1940) and Manning (1972). We thank Lilly King Manning for providing the illustrations. Gonodactylus albicinctus, new species Fig. 1 Material.—COSTA RICA: BAHIA HERRADURA; 09°38'45’"N, 84°40'55”W; 17 m; side of outer reef; mud, shell, rocks, little if any coral, sand pockets; 10 March 1972; R/V Searcher Sta. 450; M. Reaka, leg.: 16, TL 25 mm (ho- lotype, USNM 172234), 12, TL 19 mm. ISLA DEL CANO; 08°43’15’N, 83°53'07"W; 9 m; sand, coral, rubble; 14 March 1972; R/V Searcher Sta. 471; M. Reaka, leg.: 12, TL 13 mm. Diagnosis.—Rostral plate (Fig. 1b) with anterolateral angles acute but not spiniform. Ocular scales (Fig. 1b) projecting laterally, rounded. Pleura of anterior 5 abdominal somites unarmed posterolaterally. Sixth abdominal somite with 6 carinae, swollen in male at TL 25 mm, each terminating posteriorly in sharp spine. Abdominal width—carapace length indices 788— 806. Telson (Fig. la) of oerstedii-type, length and width subequal, dorsal Carinae ornamented with tubercles. Median carina strongly arched dorsally, slender in female, broadly oval in male, with posterior median spine. Ac- cessory median carinae long, almost as long as median, fused posteriorly, terminating in 3-spined corona under apex of median carina; fused part of VOLUME 92, NUMBER 3 635 Fig. 1. Gonodactylus albicinctus, female paratype, TL 19 mm: a, Posterior part of body; b, Rostral plate and ocular scales. these carinae obliterated by inflation of carinae in male. Knob prominent, with 2 spinules. Anterior submedian carinae slender, with 1 posterior spine flanked ventrally by 1-2 spinules. Carinae of submedian teeth slender, each with 1-2 dorsal spinules. Accessory intermediate, intermediate and marginal carinae sharp, accessory intermediates each with 3 dorsal spinules. 3 pairs of marginal teeth present, submedians with movable apices, intermediates sharp, laterals blunt but distinct. 11-12 submedian and 2 intermediate den- ticles present. Uropodal exopod with 10 movable spines. Color.—In life tan, with conspicuous white band across sixth abdominal somite; meral spot is white. In preservative, thoracic and abdominal somites with irregular median dark patch, those on abdomen flanked laterally by submedian dark crescent, that in turn flanked laterally by small dark spot. Telson lacking dark spots anteriorly. Measurements.—TL of male 25 mm, of 2 females 13 and 19 mm. Other measurements, in mm, of male: rostral plate length 2.0, width 1.8; carapace length 5.2; fifth abdominal somite width 4.1; telson length 3.5, width 3.5. Remarks.—This small species resembles Gonodactylus bahiahondensis 636 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON (Avra Fig. 2. Gonodactylus costaricensis, female paratype, TL 31 mm: a, Posterior part of body; b, Rostral plate and ocular scales. Schmitt, 1940, but differs in having the ocular scales more produced later- ally, the rostral plate blunt rather than spiniform anterolaterally, and more spines on the carinae of the telson, especially the accessory medians which have 1-2 dorsal tubercles. The knob in this species is not as distinctly bilobed as in G. bahiahondensis. In life the meral spot in this species is white, whereas it is powder blue in G. bahiahondensis. The two species may occur together and can be distinguished in the field by this feature. Name.—The specific epithet is from the Latin and alludes to the trans- verse white band on the abdomen of this species in life. Gonodactylus costaricensis, new species Fig. 2 Material.—COSTA RICA: BAHIA HERRADURA; 09°38'45’"N, 84°40'55”W; 20 m; tip of outer reef; mud, shell, rocks, little if any coral, sand pockets; 9 March 1972; R/V Searcher Sta. 447; M. Reaka, leg.: 22, both TL 35 mm. ISLA DEL CANO; 08°43'15"N, 83°53'07”W; 9 m; sand, coral, rubble; 14 March 1972; R/V Searcher Sta. 471; M. Reaka, leg.: 12, TL 32 mm (holotype, USNM 172235). ISLA DEL CANO; intertidal pool, in and under coral rubble and basalt boulders; March 1972; M. Reaka, leg.: 42 , TL 25-34 mm. ISLA DEL CANO; 08°42'55"N, 85°54’W; 20 m; pinnacle at corner of island close to shore, rock outcrop; 19 March 1972; R/V Searcher Sta. 491; M. Reaka, leg.: 12, TL 30 mm. Diagnosis.—Rostral plate (Fig. 2b) with anterolateral angles sharp, spi- VOLUME 92, NUMBER 3 637 niform. Ocular scales (Fig. 2b) rather broad, squarish or subrectangular, not produced laterally. Pleura of anterior 5 abdominal somites unarmed pos- terolaterally. Sixth abdominal somite with 6 sharp carinae, each produced into posterior spine. Abdominal width—carapace length indices 750-788. Tel- son (Fig. 2a) of oerstedii-type, length and width subequal or slightly broader than long, carinae ornamented with tubercles. Median carina low, sharp, terminating posteriorly in spine. Accessory median carinae usually long, extending anteriorly past midlength of median, often ornamented posteriorly with spinule and with ridge extending toward median carina. Anterior sur- face of telson usually with spined tubercle at level of intermediate carinae of sixth abdominal somite. Knob prominent, armed with 3 or 4 spinules. Anterior submedian carinae sharp dorsally, terminating in posterior spine flanked posteriorly or ventrally by second spine or spinule, carina flanked laterally and/or mesially by low but distinct carina. Carinae of submedian teeth slender, with 2 or 3 spinules anteriorly. Accessory intermediate, in- termediate and marginal carinae sharp, accessory intermediates with 1-3, usually 2, spinules dorsally. 3 pairs of marginal teeth present, submedians with movable apices. 11-12 submedian, 2 intermediate, and no lateral den- ticles present. Outer margin of uropodal exopod with 10-11 spines. Color.—In life a speckled species ranging from scarlet to mottled brown to emerald; meral spot powder blue. In preservative, body completely covered with small dark spots, abdom- inal somites with one larger spot laterally above each pleuron. Telson with 4 black spots anteriorly. Measurements.—Females only examined, TL 25-35 mm. Other measure- ments of female holotype, TL 32 mm; rostral plate length 2.5, width 2.6; carapace length 8.0; fifth abdominal somite width 6.2; telson length 5.5, width 5.5. Remarks.—This species superficially resembles Gonodactylus laliberta- densis Schmitt, 1940, but differs in that the ocular scales are not produced laterally, the anterolateral angles of the rostral plate are sharp, almost spi- niform, and the denticles of the telson are arranged differently, those on the submedian teeth being arranged in a single row rather than in two rows. In life this species has powder blue meral spots; they are white in G. lalibertadensis. Name.—The specific epithet alludes to the apparent widespread occur- rence of this species off Costa Rica. Nannosquilla canica, new species Fig. 3 Lysiosquilla decemspinosa.—Schmitt, 1940:189, fig. 20c [part, not fig. 20a, b].—Manning, 1961:30 [part]. [Not Lysiosquilla decemspinosa Rathbun, 1910.] 638 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 3. Nannosquilla canica, female holotype, TL 16.5 mm: a, Anterior part of body; b, Sixth abdominal somite, telson and uropod; c, Telson, ventral view. Material.—COSTA RICA: ISLA DEL CANO; 08°43'15”N, 85°53'07"W; 15 m; in sand around base of pinnacle; R/V Searcher Sta. 472; M. Reaka, leg.; 22 TL 15.0-16.5 mm (larger is holotype, USNM 172236). PLAYAS BLANCAS; 3-5 fms (5-9 m) 8 February 1935; Allan Hancock Pacific Expedition, Sta. 460: 1d (damaged) TL ca. 18 mm (USNM 81684). Diagnosis.—Eye stout, cornea subglobular, set obliquely on stalk. Rostral plate (Fig. 3a) pentagonal, unarmed, apex obtuse. Dactylus of claw with 11-12 teeth. Mandibular palp absent. 4 epipods present. Basal segment of each pereiopod with posterior spine. Abdomen flattened, smooth, anterior 5 somites unarmed, sixth somite pointed posterolaterally. Telson (Fig. 3b, c) broader than long, false eave produced into 3 major obtuse prominences posteriorly, blunt median and more acute submedians, latter flanked later- ally by 2 sharp, small teeth on each side. Marginal armature of telson con- sisting of, on each side, 10 submedian denticles in transverse row, | fixed submedian tooth, and 3 submarginal lateral denticles. Uropod (Fig. 3b) with 1-2 stiff setae on inner margin and 5-6 spatulate spines on outer margin of proximal segment of uropodal exopod. Inner spine of basal prolongation of uropod much the longer. Measurements.—Male damaged; females, TL 15.0-16.5 mm. Other measurements, in mm, of female holotype, 16.5 mm long: carapace length 2.5; cornea width 0.8; rostral plate length 0.8, width 1.1; fifth abdominal somite width 2.5; telson length 1.4, width 2.1. VOLUME 92, NUMBER 3 639 Remarks.—This species superficially resembles N. decemspinosa (Rath- bun, 1910), but differs in having stouter eyes, a pentagonal rather than subrectangular rostral plate with the anterolateral margins sloping toward a blunt apex and the lateral margins very divergent, more lateral marginal teeth and denticles on the telson, and a posterior spine on the basal segment of each pereiopod. The submedian marginal teeth of the telson in the two smaller specimens from Isla del Cano appear to be fixed rather than movable. They are clearly movable in the larger specimen, a damaged male. One of the three specimens from Costa Rica assigned to Nannosquilla decemspinosa by both Schmitt (1940) and Manning (1961), that taken sub- littorally, is identifiable with N. canica rather than Rathbun’s species. The other two specimens, from Isla San Lucas, Gulf of Nicoya, are identifiable with N. decemspinosa. Name.—The specific epithet is derived from the name of the type-locality, Isla de Cano. Literature Cited Manning, Raymond B. 1961. A New Lysiosquilla (Crustacea: Stomatopoda) from the Gulf of California, with a redescription of L. decemspinosa Rathbun.—Proceedings of the Bi- ological Society of Washington 74:29-35, figs. 1-6. . 1972. Stomatopod Crustacea. Eastern Pacific Expeditions of the New York Zoological Society.—Zoologica 56:95—-113, figs. 1-3. Schmitt, Waldo L. 1940. The stomatopods of the west coast of America based on collections made by the Allan Hancock Expeditions, 1933—38.—Allan Hancock Pacific Expeditions 5(4): 129-225, figs. 1-33. (RBM) Department of Invertebrate Zoology, National Museum of Natural History, Smithsonian Institution, Washington, DC 20560; (MLR) Depart- ment of Zoology, University of Maryland, College Park, MD 20740. PROC. BIOL. SOC. WASH. 92(3), 1979, pp. 640-649 STUDIES OF NEOTROPICAL CADDISFLIES XXIII: NEW GENERA FROM THE CHILEAN REGION Oliver S. Flint, Jr. Abstract.—Schajovskoya neuquenensis, n. gen., n. sp. and Androchore- ma chilense, n. gen., n. sp., belonging to the family Rhyacophilidae (Hy- drobiosinae) from Argentina and Chile, respectively, are described. The new genus Alloecentrellodes with the new species obliquus and elongatus is described from Chile. The genus is placed in the Helicophidae, to which the Chilean genera Austrocentrus and Microthremma are also transferred, thereby establishing the presence of this family in the New World for the first time. All new genera and species are described, figured, and notes on habitat given. As a result of several recent collecting trips into Chile and Argentina, a number of interesting undescribed caddisflies have been discovered. This opportunity is taken to make known those that represent undescribed genera as well as species. The first two genera are clearly representatives of the family Rhyaco- philidae, subfamily Hydrobisinae. The genus Schajovskoya (named in mem- ory of Don Sergio Schajovskoy, who was so very helpful to us in our stay at San Martin de los Andes) is a very close relative of the Chilean Para- chorema. The second rhyacophilid is Androchorema, but its relationship is very unclear although it has affinities to Rheochorema which is known from Chile and Argentina. The last, and most interesting new genus is Alloecentrellodes. This genus and several others from Chile are very closely related to certain Australian- New Zealand genera currently placed in the Helicophidae, although previ- ously placed in the Beraeidae. I have seen a few larvae from Chile that bear a strong resemblance to beraeids. Until these can be associated, described and their intimate relationships worked out, it seems best on the basis of adult characteristics to place these genera in the Helicophidae recognizing the close relationship to the Beraeidae. This is the first record of the Heli- cophidae from the New World. The females and larvae of Schajovskoya are described herein, however, these stages are unknown for the other two genera. VOLUME 92, NUMBER 3 641 Family Rhyacophilidae Schajovskoya, new genus Adult.—Spurs: 0, 4, 4. Forewing with R,,, and R,,; very short, branched at same level; M,,. and M;,, branched midway to wing margin. Hindwing with R,,, apparently arising from R,. Male abdomen without internal sacs or processes from the fifth sternum; sixth and seventh sterna with terete posteromesal processes. Male genitalia: Tenth tergum long, nearly terete, sclerotized basolaterally; paracercus platelike, filicercus long, bearing ba- sodorsally the cercus; clasper two-segmented with second segment borne dorsomesally; aedeagus tubular basally, bearing a long slender central struc- ture, divided apically into 2 asymmetrical processes. Female genitalia lat- erally compressed into a rather thin elongate structure. Larva.—Head elongate, with dark muscle scars on a pale background. Mandibles with dorsal and ventral carinae, each bearing teeth. Prosternum lacking sclerite. Foreleg with femur elongate, apicoventral process long, narrow; tibia and tarsus fused; claw with a very small basal seta. Mid and hindlegs similar, rather short, with segments broad and subequal in length; claw with basal seta very large, and angled. Lateral sclerite of anal proleg about twice as long as broad; claw with basoventral seta very large and angled. Type-species.—Schajovskoya neuquenensis, new species. The genus seems in most characteristics to be related to Parachorema Schmid. It differs in the adult in the very short R,,3; and in lacking the reflexed costal cell in the forewing. The genitalia of the two genera are also similar, but the form of the aedeagus is radically different. The larvae of Parachorema have not been described, nor is adequate material available to present diagnostic differences at this time. Schajovskoya neuquenensis, new species Figs. 1-10 This is the sole species presently known in the genus. In related genera of the family, specific diagnostic characters have been found in the male genitalia, especially the claspers, paracercus and aedeagus. Adult.—Length of forewing, 9-10 mm. Color pale brown, forewings mot- tled with paler and darker brown markings. Process of sixth sternum about 4% length of sternum, that of seventh sternum 4 length of sternum. Male genitalia: Ninth segment vertical, anterior and posterior margins parallel. Tenth tergum long, slender, sclerotized basolaterally. Paracercus broad basally with a short basoventral lobe, apex evenly rounded. Filicercus long and membranous bearing basodorsally an elongate membranous cercus. 642 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON t 4 Figs. 1-4. Schajovskoya neuquenensis: 1, male genitalia, lateral; 2, clasper, ventral; 3, female genitalia, lateral; 4, wings. VOLUME 92, NUMBER 3 643 Clasper with basal segment long, apex narrowly produced, mesally bearing short pointed setae; second segment borne dorsally at midlength of basal segment, apex hooked mesally. Aedeagus connected to clasper base by a long slender filament; bearing a long, slender central strand which divides subapically into a pair of asymmetrical, pointed processes. Female genitalia: Sixth sternum with a terete posteromesal process; seventh sternum with a minute nail-like lobe. Terminal segments long, compressed, with a complex internal structure. | Larva.—Length of prepupa, 7 mm. Head pale yellowish, darkened pos- teriorly between dark brown muscle scars, dark color extending anteriad along ecdysal lines; venter extensively darkened with a row of small, dark, muscle scars on each side posteriorly. Left mandible with ventral carina bearing 2 approximate apical teeth, dorsal carina bearing a large apicomesal tooth and a small tooth at base of central cusp; right mandible with a single large tooth at apex of both dorsal and ventral carinas. Pronotum pale yel- lowish, posterior and lateral margins black; muscle scars on postero- lateral quadrants dark brown, slightly darkened around scars. Legs pale yellow, points of articulation black. Ninth tergite pale brown. Lateral plate of anal proleg, black basally, infuscate ventrolaterally; claw golden brown. Material.—Holotype, male: Argentina, Prov. Neuquen; Arroyo Culebra, 20 km south of San Martin de los Andes, 2 Feb. 1974, O. S. Flint, Jr. USNM Type 76136. Paratypes: same data, 1¢ metamorphotype; Rio Nonthue, Estacion Forestal Pucara, 24 Feb. 1978, C. M. & O. S. Flint, Jr., 1d; Rio Aseret, near Lago Currhue, 23 Jan. 1974, O. S. Flint, Jr., 12, 16 metamorphotype. Prov. Chubut; 8 km south Hoyo de Epuyen, 10 Feb. 1974, O. S. Flint, Jr., 1d, 12 metamorphotypes. Other: Same data as holotype, | prepupa, 5 pupae; same data as Rio Aseret, 1 pupa. Habitat.—The immature stages, actually all pupae or prepupae, have been taken on 3 occasions. Both the Rio Aseret and Arroyo Culebra (as well as the Rio Nonthue where a male was swept) are virtually identical in gross appearance (Fig. 22). These streams are about 5 to 10 meters wide, by 3—5 decimeters in depth except in deeper pools. The bottom is boulders, rubble, gravel, and sand with some rocks overgrown by moss. The water is cool and clear, fast flowing, although not of an exceptional gradient. The stream 8 km south of Hoyo de Epuyen was smaller, only 2-3 meters wide by a decimeter or two deep, and the stones had a slimy coating, otherwise it was similar to the other localities. Androchorema, new genus Adult.—Spurs: 2, 4, 4. Forewing with R,,, and R,,; very short, branched at nearly same level; crossveins present (discoidal cell present); M,,,. very long, branched midway to wing margin, M,,, short, branched soon after 644 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON origin; with 2 m-cu crossveins. Hindwing with radial and medial systems disrupted by the presence of an elongate androconial structure near the anterior margin at midlength; with a hair pencil from costal margin just basad of androconia. Male lacking all process from the abdominal sterna. Male genitalia: Tenth tergum a simple, terete, membranous lobe. Cercus button-like, filicercus elongate, paracercus elongate. Clasper two-segment- ed, second segment small and poorly set off. Aedeagus with a long, spine- like ventral lip and a simple mesal process. Type-species.—Androchorema chilense, new species. The relationship of this genus is rather unclear. The forewing in its ve- nation is nearly identical to that of Rheochorema, while the aberrant struc- ture of the hindwing with its androconia 1s unique. The form of the genitalia, however, seems radically different in the two genera. The genitalia are more readily homologized with Stenochorema, but this genus lacks the second clasper segment, and has aradically different venation. At this point it seems safest to relate the genus to Rheochorema. The female and immature stages are unknown. Androchorema chilense, new species Figs. 11-13 As this is the sole species known in the genus, it is not known what the specific characteristics are, but they may be looked for in the male genitalia especially. Adult.—Length of forewing, 10 mm. Color dark brown, legs pale brown; forewing mostly denuded, membrane dark brown. Male genitalia: Ninth segment with anterior and posterior margins nearly parallel. Tenth tergum, membranous, elongate. Paracercus clavate, decumbent. Filicercus elongate, slightly curved. Cercus button-like. Clasper with basal segment elongate, broad basally, tapering apicad, inner face basally heavily sclerotized, and bearing a pair of small lobes submesally, with a distinct mesal shelf-like expansion ending in a short, finger-like lobe; apical segment small, poorly set off from apex of basal segment, terete. Aedeagus with a long, slender ventral lobe curved between bases of claspers, and a single, decurved, mesal, rod-like process. Material.—Holotype, male: Chile, Prov. Osorno; Pucatrihue, 26—30 Jan. 1978, C. M. & O. S. Flint, Jr. USNM Type 76137. Habitat.—The unique specimen was undoubtedly captured at an ultra- — Figs. 5-13. S$. neuquenensis: 5, larval head and pronotum, dorsal; 6, larval foreleg, anterior; 7, same, posterior; 8, larval midleg, posterior; 9, larval mandibles, ventral; 10, larval anal proleg, lateral. Androchorema chilense: 11, wings; 12, male genitalia, lateral; 13, same, ventral. 645 VOLUME 92, NUMBER 3 646 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON violet light. The light was close to a small brook, about a meter wide, clear and cold, with bottom of sand, gravel and rubble. Pucatrihue is on the coast of Osorno and still retains an area of Nothofagus forest, from which the stream issues. Family Helicophidae Alloecentrellodes, new genus Adult.—Ocelli lacking. Maxillary palpus short, of 4 short, subequal seg- ments; labial palpus subequal in length, of 2 segments. Head, dorsally with small posterolateral, elongate lateral, and small anterior pairs of setal warts. Pronotum with a pair of dorsal setal warts. Mesoscutum lacking setal warts, with a small anteromesal crease; scutellum with warts broadly fused across midline. Spurs: 2, 2, 4. Forewing with discoidal cell closed; R; appearing to arise from M,,., stem of M partially obsolescent; lacking 1A and 2A, with a large open cell in this area. Hindwing with R, with 3 branches, M and Cu unbranched. Type-species.—Alloecentrellodes obliquus, new species. This genus appears to be related to the Chilean Austrocentrus Schmid and even more closely to the New Zealand Alloecentrella Wise. From Aus- trocentrus it may be distinguished by the closed discal cell, presence of R,, and the large open cell in the anal area in the forewing, and the loss of an apical branch of M or Cu in the hindwing. The male of Alloecentrella has a very similar venation in the forewing but appears to have lost the stem of M or possibly Cu, and has a very different venation in the hindwing. The obvious very close relationship to the genus Alloecentrella from New Zealand, and the only slightly more distant relationship to the Australian Alloecella opens the question of proper family placement. The Australian- New Zealand component is presently placed in the family Helicophidae but was previously placed in the Beraeidae. It is certain that the Chilean com- ponent does not belong to the Sericostomatidae where it is presently placed, but in the taxon containing Alloecentrella. Therefore, Alloecentrellodes, Austrocentrus, and Microthremma are all transferred to the Helicophidae, marking the presence of the family in the New World for the first time. The relationship between the Beraeidae and Helicophidae may have to be reas- sesed at a later date when more data on the entire life cycle are known. The females and immature stages of Alloecentrellodes are unknown. Alloecentrellodes obliquus, new species Figs. 14-18 This and the following, A. elongatus n. sp., are closely related. The ninth segment is more oblique, the lateral arms of the tenth tergum surpass the tip of the cercus, and the claspers lack the ventromesal angle in A. obliquus. VOLUME 92, NUMBER 3 647 18 Figs. 14-21. Alloecentrellodes obliquus: 14, wings; 15, head and thorax, dorsal; 16, male genitalia, lateral; 17, clasper, ventral; 18, ninth and tenth terga, dorsal. A. elongatus: 19, clasper, ventral; 20, ninth and tenth terga, dorsal; 21, male genitalia, lateral. 648 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 22. Arroyo Culebra, type-locality of S. neuquenensis. Adult.—Length of forewing, 7 mm. Color fuscous, apices of legs very slightly paler; forewing fuscous, immaculate. Seventh sternum with a broad, nail-like process posteromesally. Male genitalia: Ninth segment very oblique. Tenth tergum tripartite: a long, slender dorsomesal process; paired lateral processes clearly surpassing apex of cercus and ending in a short spine. Cercus elongate. Clasper with basal segment elongate, tapering, apicomesal face rugose; apical segment only partially distinct from basal segment, elon- gate, hooked mesad. Aedeagus long, slender, with slender lateral flanges and a dark apicomesal structure. Material.—Holotype, male: Chile, Prov. Malleco, Parque Nacional Con- tulmo (near boundary of Prov. Arauco), 19 Oct. 1969, Flint & Barria. USNM Type 76138. Paratype: Cordillera Nahuelbuta, Cabreria, 1100 m., 9-15 Jan. Oi ee eena Guy lc Habitat.—My notes for 19 October indicate that adult caddisflies, pre- sumably including the holotype, were swept from small streams just beyond the Arauco Provincial boundary. The small streams crossing the road here flow steeply down the forested hillsides, and are barely a half-meter wide by a decimeter deep. The waters are clear and cold with a substrate of sand, gravel and rubble. VOLUME 92, NUMBER 3 649 Alloecentrellodes elongatus, new species Figs. 19-21 This species is closely related to the previous species. The ninth segment is horizontal, the lateral arms of the tenth tergum do not surpass the cercus, and the claspers have a distinct ventromesal angle in A. elongatus. Adult.—Length of forewing, 5 mm. Color fuscous, legs slightly paler; forewing fuscous, immaculate. Seventh sternum with a broad, nail-like pro- cess posteromesally. Male genitalia: Ninth segment horizontal. Tenth ter- gum tripartite: a long, slender, dorsomesal process; paired lateral processes not attaining apex of cercus, and ending in a short spine. Cercus elongate. Clasper with basal segment elongate, produced into short lobes ventrome- sally; apical segment only partially distinct from basal segment, elongate, ending bluntly, slightly angled mesad. Aedeagus long, slender, with well- developed lateral flanges and a dark mesal structure. Material.—Holotype, male: Chile, Prov. Nuble; Recinto, 14 Oct. 1969, Flint & Barria. USNM Type 76139. Paratype: Same data, 16. Habitat.—My notes indicate that 2 adult caddisflies, of necessity the two listed above, were collected from a small hillside streamlet. The habitat is essentially the same as that of A. obliquus, however, the surroundings are not forested but densely overgrown by tall brush. Department of Entomology, Smithsonian Institution, Washington, D.C. 20560. PROC. BIOL. SOC. WASH. 92(3), 1979, pp. 650-657 CARPIAS RICHARDSON 1902, A SENIOR SYNONYM OF BAGATUS NOBILI 1906, AND THE VALIDITY OF CARPIAS MINUTUS (RICHARDSON 1902) (ISOPODA: ASELLOTA: JANIRIDAE) Thomas E. Bowman and Byron F. Morris Abstract.—lIllustrations and descriptions of Carpias bermudensis Rich- ardson and Janira minuta Richardson are offered as evidence that these species are distinct and not synonymous as claimed by Menzies and Glynn (1968). However, their opinion that Carpias Richardson and Bagatus Nobili are synonyms appears to be justified, and the genus takes the older name Carpias. The genus Carpias was proposed by Richardson (1902) for a small asellote isopod of the family Janiridae from Bermuda, characterized by having ex- traordinarily large gnathopods in the male, longer than the entire body. The 1902 description and figures of C. bermudensis were repeated in her 1905 monograph. No new records appeared until 1968, when Miller (1968) re- ported it from Florida. The same year Menzies and Glynn (1968) reported its occurrence in the vicinity of La Parguera, Puerto Rico, and transferred it to the genus Bagatus Nobili (1906). Another Bermudan asellote, Janira minuta Richardson (1902), which had been transferred to Bagatus by Nor- denstam (1946), was declared by Menzies and Glynn to be a synonym of Bagatus bermudensis. Nobili (1906, 1907) made no mention of Carpias when he proposed Ba- gatus, and Richardson’s (1902) work on Bermudan isopods is not cited in either of Nobili’s papers. Either Nobili was unaware of Carpias, or consid- ered it generically distinct from Bagatus. Inaccuracies in Richardson’s de- scription have puzzled others; Wolff (1962:45) noted that ‘‘the mandible seems to be totally without molar process according to fig. 505a, but Miss Richardson does not mention this feature in her description.’’ In fact, C. bermudensis has a well developed molar. In his key to genera of Janiridae, Menzies (1962) separated Carpias from Bagatus by the presence of a dactyl on pereopod 1 in Carpias and its absence in Bagatus. In the type-species of Bagatus, B. stylodactylus, the — Fig. 1. Carpias minutus: a, 8 dorsal; b, ¢ dorsal; c, o lateral; d, 2 head; e, 2 right uroped, dorsal. VOLUME 92, NUMBER 3 652 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON f ff y Uf TIAN, ny Fig. 2. Carpias minutus °: a, Left mandible; b, Maxilla 1; c, Maxilla 2; d, Maxilliped; e, Maxilliped, distal segment of palp. VOLUME 92, NUMBER 3 653 m | k l o.1 mm, n 0 figs. e-o Fig. 3. Carpias minutus: a, Pereopod 1, 2; b, Spine from carpus of same; c, Pereopod 1, 3; d, Dactyl and distal end of propus of same; e-i, Dactyls of 2 pereopods 2, 3, 4, 6, 7; j-o, Dactyls of ¢ pereopods 2-7. 654 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 0.4 mm —— ee Fig. 4. Carpias minutus 3: a-d, Pleopods 1-4. Carpias bermudensis 3: e, Head and pe- reonite 1, dorsal; f, Right mandible; g, Incisor and lacinia of left mandible; h, Maxilla 1; i, Maxilla 2. VOLUME 92, NUMBER 3 655 Fig. 5. Carpias bermudensis: a, Maxilliped; b, Pereopod 1, adult 3; c, Pereopod 1, juvenile 3 (2 mm), setae omitted; d, Pereopod 1, 2; e, Pereopod 7, 2; f, Dactyl of same; g, Pleopod 1, 6; h, Pleopod 2, 6. 656 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON dactyl is reduced to a small knob, but in other species the dactyl is well developed as in Carpias. No other differences between C. bermudensis and Bagatus spp. have generic value. The enormous 6 pereopod | of the former is quantitatively rather than qualitatively different from this pereopod in ¢ Bagatus. Menzies and Glynn were quite justified in fusing the 2 genera. However, they erred in using the younger name, Bagatus, rather than the older Carpias. All species of Bagatus, the 9 listed by Monod (1961) and B. serricaudus Men- zies and Glynn (1968) must be transferred to Carpias. Janira falcifera Bar- nard (1962) has a very large 6 pereopod 1; hence it appears to be a Carpias. While we agree that Carpias and Bagatus are congeneric, we do not accept Menzies and Glynn’s (1968) proposition that C. bermudensis and C. minutus are the same species. We discuss below differences we have found between C. minutus, based on specimens taken from Sargassum off Ber- muda, in which it is common (Morris and Mogelberg, 1973), now deposited in the National Museum of Natural History, and syntypes of C. bermudensis (USNM 24865). The latter are in poor condition, but enough details could be discerned to confirm the distinctness of the 2 species and to give the comparison that follows. The lateral margins of pereonite 1 are incised in C. bermudensis, but not in C. minutus. The mouthparts are similar, with minor differences in seta- tion. Mandibular palp segment 3 is longer and narrower in C. minutus; the incisors and lacinia are 5-cuspate in both species. The 3 pereopod 1 shows great differences in both young and fully mature specimens. The merus is very elongate in C. bermudensis. The carpus is deltoid in C. minutus, elongate-mitten-shaped in C. bermudensis; its palm is transverse in C. minutus, oblique in C. bermudensis, and has 3 teeth in C. minutus (including the long distal tooth), 2 in C. bermudensis. The pro- pus of C. minutus is toothless and has parallel margins; that of C. bermu- densis has 2 teeth and widens distally into a bulbous apex. The dactyls of both species are biungulate in pereopod 1, triungulate in pereopods 2-7. The ¢ pleopod 1 has stronger outer lobes in C. minutus. Inner lobes are developed in C. minutus, not in C. bermudensis. The 6 pleopod 1 is a most important character in Janiridae taxonomy, and one would be justified in separating C. bermudensis from C. minutus on this character alone. Literature Cited Barnard, K. H. 1962. New records of marine Crustacea from the East African region.— Crustaceana 3(3):239-245. Menzies, Robert J. 1962. The zoogeography, ecology, and systematics of the Chilean marine isopods. Reports of the Lund University Chile Expedition 1948-49:42.—Lunds Uni- versitets Arsskrift. N.F. Avd. 2, STAD) 1=162: , and Peter W. Glynn. 1968. The common marine isopod Crustacea of Puerto Rico.— Studies on the Fauna of Curacao and other Caribbean Islands 27(104): 1-133. VOLUME 92, NUMBER 3 657 Miller, Milton A. 1968. Isopoda and Tanaidacea from buoys in coastal waters of the conti- nental United States, Hawaii, and the Bahamas (Crustacea).—Proceedings of the United States National Museum 125(3652): 1-53. Monod, Théodore. 1961. Sur un isopode asellote du genre Bagatus recueilli sur un poisson du Sénégal.—Crustaceana 2(1):68-77. Morris, Byron F., and Deborah D. Mogelberg. 1973. Identification manual to the pelagic Sargassum fauna.—Bermuda Biological Station for Research Special Publication 11: 1-63. Nobili, Giuseppe. 1906. Diagnoses préliminaires de Crustacés, Décapodes et Isopodes nou- veaux recueillis par M. le Dr. G. Seurat aux [les Touamotou.—Bulletin du Muséum d’Histoire Naturelle 1906, no. 5:256—270. . 1907. Ricerche sui crostacei della Polinesia. Decapodi, stomatopodi, anisopodi e iso- podi.—Memorie della Reale Accademia delle Scienze di Torino, ser. 2, 57:351—430, pls. 1-3. Nordenstam, Ake. 1946. Marine Isopoda from Professor Dr. Sixten Bock’s Pacific Expedition 1917—1918.—Arkiv for Zoologi 37A(7): 1-31. Richardson, Harriet. 1902. The marine and terrestrial isopods of the Bermudas, with descrip- tions of new genera and species.—Transactions of the Connecticut Academy of Sciences 11:277-310, pls. 37-40. . 1905. A monograph on the isopods of North America.—Bulletin of the United States National Museum 54:I—-LIII + 1-727. Wolff, Torben. 1962. The systematics and biology of bathyal and abyssal Isopoda Asellota.— Galathea Report 6:1—320, pls. 1-19. (TEB) Department of Invertebrate Zoology, Smithsonian Institution, Washington, DC 20560; (BFM) Outer Continental Shelf Office, Bureau of Land Management, 555 Cordova St., Box 1159, Anchorage, AK 99501. PROC. BIOL. SOC. WASH. 92(3), 1979, pp. 658-664 REDESCRIPTION OF HALIOPHASMA BARNARDI (MONOD) FROM WEST AFRICA (CRUSTACEA: ISOPODA: ANTHURIDAE) Brian Kensley Abstract.—Haliophasma barnardi (formerly in the genus Notanthura Monod) is recorded from a mesohaline creek in the Niger delta, Nigeria. The female is redescribed and figured, and its position within Haliophasma briefly discussed. Three anthurids were collected during 1978-79 in the Niger delta by Dr. C. B. Powell of the School of Biological Sciences, University of Port Har- court, Nigeria, and submitted to the Smithsonian for identification. The specimens proved to be a species first described in 1927 and not recorded since. This is also the first time the whole animal has been seen, the original material having come from fish stomach contents. The present paper is one of a series of short contributions dealing with the taxonomy of the anthurid isopods, preliminary to a major generic revi- sion of the group. Family Anthuridae Haliophasma barnardi (Monod) Figs. 1-3 Notanthura barnardi Monod, 1927:202, figs. 1—9.—Nierstrasz, 1941:239.— Poore, 1975:532. Description.—@ . Integument hardly indurate. Body proportions: C < 1 > 2>3<4=5>6*>7. Cephalon with low rostrum, not reaching beyond anterolateral corners; eyes lacking. Pereonites 3—6 with shallow middorsal pit. Pleonites 1-5 fused, indicated laterally by short slits; pleonite 6 free, with middorsal notch in posterior margin. Telson dorsally gently convex, widest at midlength, tapering to broadly rounded apex; with 2 proximal Statocysts. Antennular peduncle with basal segment equal to 3 distal segments to- gether; segment 3 with 3 elongate simple setae; segment 4 short, with single simple seta and sensory seta; flagellum of 4 articles, 3 distal articles very short, basal article elongate, with 2 distal aesthetascs, 3 distal articles each with single aesthetasc. Antennal peduncle 5-segmented, segment 2 grooved to accommodate antennule, with hair-like setae on medial margin; segments 3 and 4 subequal; segment 5 slightly longer; flagellum of 4 articles, each VOLUME 92, NUMBER 3 659 Fig. 1. Haliophasma barnardi: a, Female in dorsal view; b, Antennule; c, Antenna; d, Mandible; e, Maxilla; f, Maxilliped. with distal cluster of simple setae. Mandibular palp 3-segmented, middle segment somewhat longer than basal segment; basal segment with single elongate simple seta; segment 2 with 5 elongate setae; terminal segment with 3 distal fringed spines and clusters of short setules; incisor broad, strongly sclerotized, indistinctly cusped; lacinia with 13 marginal teeth; mo- lar acutely triangular, with row of spines on upper surface. Maxilla with 1 strong sclerotized spine and 7 smaller distal spines. Maxilliped 4-segmented, 660 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 2. Haliophasma barnardi: a, Pereopod 1; b, Pereopod 2; ¢, Pereopod 7; d, Pleopod 1. VOLUME 92, NUMBER 3 661 ‘ Wi Mf 7 | es ee ¢ ; f tha Yad ty ZN Fig. 3. Haliophasma barnardi: a, Pleon in lateral view; b, Telson and uropods in dorsal view. 662 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON with possible trace of incomplete suture on segment 3; segment 2 with fine hair-like setae on outer surface; segment 3 with 4 short setae on medial margin, few scattered and more elongate setae on outer surface; segment 4 short, semicircular, with 5 setae. Pereopod | subchelate, robust; strongly sclerotized unguis about half length of dactylus, with short stout basal ac- cessory spine; propodus proximally expanded, palm with slight lobe/notch at about midpoint and several marginal setae, 6 more elongate setae on inner surface near dactylar articulation; carpus triangular, with several simple setae on posterior margin and inner surface, few fringed scales at rounded apex. Pereopods 2 and 3 less robust than pereopod 1, not subchelate; unguis about one-third length of dactylus; propodus proximally only slightly wider than distally, with elongate simple setae on anterior and posterior margins; strong serrate spine at posterodistal corner; carpus short, triangular, distally broadly rounded, with elongate simple setae on posterior margin; merus with elongate simple setae on both margins; ischium with elongate simple setae on posterior margin only. Pereopods 4—7 unguis about one-third length of dactylus; propodus parallel-sided, narrow, with row of fringed scales on posterior margin, strong serrate spine at posterodistal corner; carpus elon- gate rectangular, with row of elongate simple setae on posterior margin, sensory spine at posterodistal corner, 3 elongate setae on anterodistal mar- gin; merus with elongate simple setae on both margins. Pleopod 1 exopod operculiform, with strong groove on outer surface, numerous plumose setae on distal margin; endopod about half width of and slightly shorter than exopod, distal quarter constricted, apically rounded, with 8 plumose setae; basis with 4 retinaculae. Uropodal exopod not quite reaching endopod base, roughly oval, outer margin sinuous, with dense fringe of plumose setae, apically acute; endopod triangular, apically rounded, reaching telsonic apex. Material examined.—Paris Museum, Syntypic material: numerous frag- ments and appendages, including 7 more or less complete heads and varying numbers of attached pereonites; taken from stomach of Trygon margarita, Souelaba, Cameroun, West Africa, Nov. 25, 1925. British Museum, BM.1928.9.25.1—2. Syntypic material: 3 heads with vary- ing numbers of pereonites and appendages; Souelaba, Cameroun, West Af- rica. USNM 172266-8, 32, TL 6.8 mm, 7.0 mm, 9.1 mm; from Niger delta near Port Harcourt, Nigeria; taken from bed of leaf fragments and sticks Over anoxic mud in mesohaline mangrove creek; collected Aug. and Nov. 1978, Feb. 1979. Remarks.—There is no doubt that the present material from the Niger delta is the same species which Monod described from Souelaba, Bay of Douala, Cameroun (about 300 kilometers to the southeast). Monod’s ma- terial came from the stomach of the stingray Trygon margarita which is known to inhabit mangrove estuaries (D. Cohen, pers. comm.). The general VOLUME 92, NUMBER 3 663 morphology of the body and appendages of the Nigerian and Cameroun material agree well. A few differences between Monod’s figures and the present specimens may be noted. Monod shows a relatively broader and squatter maxilliped, 5 distal maxillar spines, and does not show a row of fine teeth on the acutely triangular molar. However, a mounted mandible, maxilla, and maxilliped taken from the Paris Museum material agree exactly with the Nigerian specimens. Further, the structure of the antennular fla- gellum agrees even to the number of aesthetascs per article. Poore (1975) redefined the genus Haliophasma from which the present material differs on two points, viz. the lack of eyes, and a 4-articulate, rather than a 2-articulate antennular flagellum. Kensley (1975) described Halio- phasma caeca from the west coast of South Africa, and noted the lack of eyes in that species. The figure of the maxilliped (Fig. 1G) shows a 3-seg- mented appendage; in fact, the suture of the terminal setose segment was not indicated, and the maxilliped of H. caeca is remarkably similar to that of H. barnardi. The overall similarity between H. caeca and H. barnardi is striking, especially in the setose pereopods 2-7. This, with the lack of eyes, is probably an adaptation to habitat; in both cases the animals seem to be part of the infauna of organically rich sandy or muddy substrates. The only feature of H. barnardi which gives pause is the dentate and acutely triangular mandibular molar. Acutely triangular molars are found in e.g. H. cycneum Poore, H. cribensis Poore, and H. pinnatum Poore, but the fine dentition of the present molar does not seem to have been noted previously. Whether this feature together with the lack of eyes and the 4-articulate antennular flagellum are sufficient to separate Notanthura from Haliophas- ma is a moot point. In view of the many points of agreement, however, the two blind species (H. barnardi and H. caeca) could perhaps be regarded as forming a subgenus within Haliophasma. The monotypic genus Notanthura Monod is thus synonymized with Haliophasma Haswell. The only other known species of Haliophasma from West Africa is H. dakarensis Barnard, 1925. This is a much larger indurate species, possessing eyes and a distinctly keeled telson. Acknowledgments I am grateful to Dr. C. B. Powell of the School of Biological Sciences, University of Port Harcourt, Nigeria, for donating the three anthurid spec- imens to the Smithsonian Institution, and for supplying valuable collection data; and to Miss J. Ellis of the British Museum (Natural History) and Dr. J. Forest of the Paris Museum, for the loan of type-material. I wish to thank Miss Mary-Jacque Mann of the Smithsonian Scanning Electron Microscope Laboratory for assistance with the micrographs, and Dr. Thomas E. Bow- man of the Smithsonian Institution, for his critical reading of the MS. 664 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Literature Cited Barnard, K. H. 1925. A revision of the family Anthuridae (Crustacea Isopoda) with remarks on certain morphological peculiarities.—Journal of the Linnaean Society, London (Zo- ology) 36: 109-160. Kensley, B. 1975. A new species of anthurid isopod from the Cape.—Zoologica Africana 10:209-213. Monod, T. 1927. Notes isopodologiques II. Sur un Anthuride nouveau du Cameroun Notan- thura barnardi nov. gen., nov. sp.—Bulletin de Société Zoologique de France 52:200- Dlale Nierstrasz, H. F. 1941. Die Isopoden der Siboga-Expedition IV. Isopoda Genuina III. Gna- thiidea, Anthuridea, Valvifera, Asellota, Phreatocoidea.—Siboga Expedition monograph 32d:235-306. Poore, G. C. B. 1975. Australian species of Haliophasma (Crustacea: Isopoda: Anthuridae).— Records of the Australian Museum 29:503-533. Department of Invertebrate Zoology, Smithsonian Institution, Washing- ton, D.C. 20560. PROC. BIOL. SOC. WASH. 92(3), 1979, pp. 665-670 REDESCRIPTION OF ZONOPHRYXUS TRILOBUS RICHARDSON, WITH NOTES ON THE MALE AND DEVELOPMENTAL STAGES (CRUSTACEA: ISOPODA: DAJIDAE) Brian Kensley Abstract.—The mature female holotype of the dajid isopod parasite Zo- nophryzus trilobus Richardson is redescribed. Cryptoniscid larvae, males, and immature females from a specimen of pandalid shrimp Heterocarpus gibbosus, taken from the Philippines, are suggested to be conspecific with Z. trilobus and are described. It is also suggested that the immature female represents a possible evolutionary stage between the Bopyridae and the Dayjidae. While identifying caridean decapods collected by the R/V Albatross in Philippine waters in 1909, Dr. F. A. Chace, Jr., drew my attention to three small parasitic isopods on the carapace of a pandalid shrimp. After exami- nation of these isopods, I decided that these possibly represented an im- mature phase of a previously described dajid. This paper provides descrip- tions of these newly-found cryptoniscid larvae, the male, and the immature female, as well as the mature female of Zonophryxus trilobus, and discusses the possible evolutionary implications of these developmental stages. Suborder Epicaridea Family Dajidae Zonophryxus trilobus Richardson Figs. 1—4 Zonophryxus trilobus Richardson, 1910:41, fig. 39.—Barnard, 1914:230.— Nierstrasz and Brender a Brandis, 1923:111.—Holthuis, 1949:7. Description.—Cryptoniscid Larva: (Fig. la). Body dorsally convex, ta- pering posteriorly. Cephalon with semicircular anterior margin. Eyes ab- sent. Pereon of 7 free segments; coxae anterolaterally rounded, postero- distally acute. Pleon of 6 free segments, decreasing in length and width posteriorly; last segment broadly triangular, apex evenly rounded. Anten- nules medially almost contiguous; basal segment attached transversely, with few setae; 2nd segment tripartite, consisting of broad proximal part, narrow lobe bearing 2 terminal setae, and low rounded tubercle bearing several 666 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 05 Fig. 1. Zonophryxus trilobus: a, Cryptoniscid larva in ventral view; b, Male in ventral view. aesthetascs; terminal segment relatively elongate, bearing few terminal se- tae. Antenna elongate, of 4 relatively elongate proximal segments, and few distal articles (some possibly missing). Oral cone with anterior shield-like labium; 2 hollowed, distally sclerotized mandibles projecting from oral tube. 7 pairs of subcheltate pereopods present, becoming slightly more elongate posteriorly; propodal palm bearing 2 serrate spines; carpus triangular, with distal spine becoming serrate in posterior pereopods. 5 pairs pleopods pres- ent, becoming shorter and narrower posteriorly; each with broadly trian- gular protopodite and 2 rami each bearing several elongate simple setae distally. Uropods with stout protopodite equal in length to cylindrical rami; VOLUME 92, NUMBER 3 667 Fig. 2. Zonophryxus trilobus: Ventral and dorsal view of immature female. inner ramus slightly wider than outer; both with 2 short distal spines and single elongate spine. Male.—(Fig. 1b). Cephalon and Ist pereonite fused; cephalothorax oval in outline. Eyes absent. 6 free laterally-rounded pereonites present. Pleo- telson a single pyriform segment. Antennule a single briad segment bearing few short spines, with single setae on terminal papilla. Antenna of 4 rela- tively elongate basal segments and ?2 distal articles. Hollowed sclerotised mandibles projecting from oral cone. 7 pairs robust subchelate pereopods present, each with concave propodal palm bearing proximal spiniform pro- cess. Pleotelson lacking appendages except for small pair uniramous uro- pods situated in ventrodistal hollow. Immature female.—(Fig. 2). Body dorsally strongly convex, oval in out- line; cephalon, pereonites and pleonites fused, segmentation of pereon and pleon indicated by notches and small points on circumference; faint indi- cations of segmentation middorsally. Cephalon with broad convex flange. Pereonite | fused with cephalon. Antennule expanded into lamella embrac- ing oral cone. Antenna a simple digitiform process attached to outer region of antennular lamella. Oral cone with 2 projecting sclerotized mandibles. Median sternum posterior to oral cone raised into ridge embraced by large bilobed oostegite of pereopod 1. 6 pairs subchelate pereopods present, each of 6 segments (including coxa); propodal palm slightly convex. Pereopods 668 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 3. Zonophryxus trilobus: Ventral and dorsal view of mature female holotype. 2-5 with small oval oostegites; pereopod 6 lacking oostegite; body wall between pereopods 5 and 6 with small lamellar fold. Pleonal segments in- dicated by curved transverse grooves in sternum. Mature female.—(Fig. 3). Body dorsally strongly convex, oval in outline, with faint irregular lines and grooves indicating segmentation. Ventral flange anteriorly wide, converging posteriorly, with several notches and points on circumference, 4 posterior points on each side close together. Antennules, antennae, and oral cone as in immature ¢. 5 pairs subchelate pereopods. Median sternum posterior to oral cone raised into ridge, embraced by oval oostegite of pereopod 2 overlying oval oostegite of pereopod 1. Oostegites of pereopods 3 and 4 broad, membranous, reaching ventral midline, covered ventrally by broad expansion of body wall posterior to pereopod 5, covering entire posterior ventral surface, meeting midventrally, divided into large anterior and smaller posterior lobe. Material.—Zonophryxus trilobus: Holotype, mature 2, USNM 40927, TL 13.0 mm. Albatross sta. 5259, off Caluya Is., Sulu Archipelago, Philippines, 570 m. Immature 9 ?, TL 5.0 mm, 4.1 mm, 3.0 mm; 2¢ 6. TL 1.4 mm; Cryptoniscid larvae, TL 1.0 mm. Albatross sta. 5622, west of Halmahera Is., Indonesia, 503 m. Habitat.—Two of the three immature females were found attached to the carapace at the left base of the rostrum of the pandalid shrimp Heterocarpus VOLUME 92, NUMBER 3 669 Fig. 4. Anterior carapace and rostral base of Heterocarpus gibbosus with puncture marks and immature female Zonophryxus trilobus in situ. gibbosus Bate, 1888 (see Fig. 4). In the same region of the carapace were several small circular punctures each surrounded by a discolored area, pre- sumably the result of the parasites’ feeding. The third female was found loose in the bottle with the shrimp. The largest immature female had a male attached to the pleon, and a cryptoniscid in the cephalon region. The small- est female had a cryptoniscid amongst the pereopods, while the third female had a male on the ventral surface. The holotype of Z. trilobus, unfortunately, was taken from a trawl free of any host. Remarks.—On the basis of several similarities, it is proposed that the small hemispherical isopods taken from the anterior carapace of Hetero- carpus gibbosus could be immature females of Zonophryxus trilobus Rich- ardson. The similarities include similar structure of the fused antennules and antennae, pereopod structure, the raised mediosternum of the anterior pereonites embraced by the oostegite of pereopod 1, the notches and points in the circumference indicating the segmentation of the pereon and pleon. It is suggested that development of the ovaries/eggs leads to a general distention of the body, especially in the region of pereonites 5 and 6 (as indicated by the increased distances between notches on the circumference flange of the mature female). This distention, being mainly in the dorsal region, leads to the convergence of the posteroventral body margin char- acteristic of Zonophryxus. At the same time, the lamellar section of the body wall between pereopods 5 and 6 increases in size, leading to a complete closure of the ventral brood pouch and a simultaneous suppression of pe- reopod 6. 670 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON The presence of cryptoniscid larvae on the small females is perhaps also evidence of immaturity. The broad lamellar outgrowths on each side of the body wall in the region of the Sth and 6th pereopods are also seen in Z. dodecapus (Holthuis, 1949) where they are referred to as oostegites. In this Atlantic species, the sixth pereopods are retained in the mature female and are situated posteriorly. Apart from the development of eggs and the brood pouch, the loss of the 6th pair of pereopods seems to be the major change from immature to mature female. This cannot be regarded as unusual, as the change from cryptoniscid larva to immature female also involves the loss of a pair of legs. The immature female form described here may represent a stage in the evolution between bopyrid and dajid epicarideans. The change from cryp- toniscid to adult female bopyrid (the latter generally with seven pairs of pereopods, five pairs of pleopods and one pair of uropods) requires little change in the state of the pereonal and pleonal appendages. The next step towards the more modified dajids is represented by the immature female described above, with six pairs of pereopods and some pleomeres distin- guishable, but no pleopods. Loss of another one or two pairs of pereopods, and thereby the concentration of the remaining pereopods in the anterior pereon, and complete obliteration of the pleomeres, would result in the more typical dajid. Acknowledgments My sincere thanks are due to Dr. F. A. Chace, Jr., of the Smithsonian Institution, Washington, both for drawing my attention to the parasites de- scribed herein, and for reading the MS; also to Dr. T. E. Bowman of the Smithsonian Institution, and Dr. J. C. Markham of the Bermuda Biological Station for Research, for their many useful comments and criticisms on the paper. Literature Cited Barnard, K. H. 1914. Contributions to the Crustacean Fauna of South Africa 1. Additions to the Marine Isopoda.—Annals of the South African Museum 10:197—230. Bate, C. S. 1888. Report on the Crustacea Macrura collected by H.M.S. Challenger during the years 1873—-1876.—Report of the Voyage of the H.M.S. Challenger 1873-1876 24: 1— 942. Holthuis, L. B. 1949. Zonophryxus dodecapus nov. spec., a remarkable species of the family Dajidae (Crustacea Isopoda) from the Canary Islands.—Proceedings of the Koninklijke Nederlandsche Akademie van Wetenschappen 52:208-213. Nierstrasz, H. F., and G. A. Brender a Brandis. 1923. Die Isopoden der Siboga-Expedition II. Isopoda Genuina I. Epicaridea.—Siboga Expedition monograph 32b:57-121. Richardson, H. 1910. Marine Isopods collected in the Philippines by the U.S. Fisheries Steam- er Albatross in 1907-8.—Bureau of Fisheries Document 736: I-44. Department of Invertebrate Zoology, Smithsonian Institution, Washing- ton, DC 20560. _ the editors. INFORMATION FOR CONTRIBUTORS Content.—The Proceedings of the Biological Society of Washington contains papers bearing on systematics in the biological sciences (both botany and zoology, including paleontology), and notices of business transacted at meetings of the Society. Except at the direction of the Council, only manuscripts by Society members will be accepted. Papers will be published in English, except for Latin diagnosis/description of plant taxa which should not be duplicated by an English translation, or summary in an alternate language when appropriate. Submission of manuscripts.—Manuscripts should be sent to the Editor, Proceedings of the Biological Society of Washington, National Museum of Natural History, Washington, D.C. 20560. Review.—One of the Society’s aims is to give its members an opportunity for prompt pub- lication of their shorter contributions. Manuscripts will be reviewed in order of receipt by a board of associate editors and appropriate referees. The Proceedings are issued four times a year. Presentation.—Clarity of presentation and requirements of taxonomic and nomenclatural procedures necessitate reasonable consistency in the organization of papers. Authors should follow recent issues of the Proceedings as models, including an abstract. Telegraphic style is recommended as the most economical of space for descriptions. Synonymy of abbreviated style (author, date, page) with full citations only in Literature Cited is also recommended. The establishment of new taxa must conform with the requirements of the appropriate in- ternational codes of nomenclature. Authors are expected to be familiar with these codes and to comply with them. New species-group accounts must designate a type-specimen deposited _ in an institutional collection. The sequence of material should be: Title, Author(s), Address(es), Abstract, Text, Literature 1 Cited, Appendix, Tables (each table numbered with an Arabic numeral and heading provided), List of Figures (entire figure legends), Figures (each numbered and identified). Type manuscripts double-spaced throughout (including tables, legends, and footnotes) on one side of paper measuring approximately 8/2 x 11 inches, leaving margins of at least one inch all around. Submit a facsimile with the original and retain an author’s copy. Number pages , consecutively at the top. One manuscript page = approximately 2 of a printed page. Underline singly scientific names of genera and lower categories; leave other indications to Figures and tables, with their legends and headings, should usually be self explanatory, not requiring reference to the text. Indicate their approximate placement by a pencil mark in the _ margin of the manuscript. Illustrations should be planned in proportions that will efficiently use space on the type bed of the Proceedings (11.8 x 18 cm). Legends require 4 mm of column length per line. Proofs.—Galley proof will be submitted to authors for correction and approval. Changes _ other than printer’s or editor’s errors may be charged to authors. Reprint orders will be taken _ with returned proof. CONTENTS Lumbrineridae, Arabellidae, and Dorvilleidae (Polychaeta), principally from Florida, with descriptions of six new species Thomas H. Perkins Four new species of Carazziella (Polychaeta: Spionidae) from North and South Amer- ica, with a redescription of two previously described forms James A. Blake The Indo-Pacific pipefish genera Notiocampus gen. nov. and Nannocampus Gunther C. E. Dawson An Old World occurrence of the Eocene avian family Primobucconidae Storrs L. Olson and Alan Feduccia A new species of Eleutherodactylus from northern Ecuador (Amphibia: Leptodac- tylidae) John D. Lynch Peosidrilus biprostatus n. g., n. sp., a marine tubificid (Oligochaeta) from the eastern United States H. R. Baker and Christer Erséus New gastropods from the Abrolhos Archipelago and reef complex, Brazil Edward J. Petuch Voorthuyseniella ventressi, new species (Microproblematica) from the basal Pleistocene of Louisiana Drew Haman A revision of the milliped genus Delophon, with the proposal of two new tribes in the subfamily Abacioninae (Callipodida: Caspiopetalidae) — Rowland M. Shelley A synopsis of the milliped genus Cambala, with a description of C. minor Bollman (Spirostreptida: Cambalidae) Rowland M. Shelley Note on the occurrence of a swarm of Salpa cylindrica Cuvier (Tunicata: Salpidae) in Sardinera Lagoon, Puerto Rico Takasi Tokioka and P. B. Bhavanarayana Louisiana and Panama Canal locations and ecology of Munna (Pangamunna nov. sub- gen.) reynoldsi Frankenberg & Menzies (Isopoda: Asellota) George A. Schultz Stenocorophium bowmani, a new genus and species of the family Corophiidae from the Palau Islands (Crustacea: Amphipoda) Gordan S. Karaman A new species of primary burrowing crayfish of the genus Cambarus from the Ridge and Valley province in Tennessee Raymond W. Bouchard and David A. Etnier A new North American species of fresh-water Tubificidae (Oligochaeta) Michael S. Loden Revision of some Polydorids (Polychaeta: Spionidae) described and recorded from British Columbia by Edith and Cyril Berkeley James A. Blake Additional notes on cave shrimps (Crustacea: Atyidae and Palaemonidae) from the Yucatan Peninsula, México H. H. Hobbs III Three new stomatopod crustaceans from the Pacific coast of Costa Rica Raymond B. Manning and Marjorie L. Reaka Studies of Neotropical caddisflies XXIII: new genera from the Chilean region Oliver S. Flint, Jr. Carpias Richardson 1902, a senior synonym of Bagatus Nobili 1906, and the validity of Carpias minutus (Richardson 1902) (Isopoda: Asellota: Janiridae) Thomas E. Bowman and Byron F. Morris Redescription of Haliophasma barnardi (Monod) from West Africa (Crustacea: Isopoda: Anthuridae) Brian Kensley Redescription of Zonophryxus trilobus Richardson, with notes on the male and devel- opmental stages (Crustacea: Isopoda: Dajidae) Brian Kensley 658 665 Proceedings of the BIOLOGICAL SOCIETY " WASHINGTON Volume 92 16 November 1979 Number 4 THE BIOLOGICAL SOCIETY OF WASHINGTON 1978-1979 Officers President: Oliver S. Flint, Jr. Secretary: Michael A. Bogan Vice President: Richard Banks Treasurer: David L. Pawson Elected Council Arthur H. Clarke W. Duane Hope Robert H. Gibbs, Jr. Catherine J. Kerby Austin B. Williams PROCEEDINGS Editor: C. W. Hart, Jr. Associate Editors Classical Languages: George C. Steyskal Invertebrates: Thomas E. Bowman Plants: David B. Lellinger Vertebrates: Richard Banks Insects: Robert D. Gordon Membership in the Society is open to anyone who wishes to join. There are no prerequisites. Annual dues of $7.00 include subscription to the Proceedings of the Biological Society of Washington. Correspondence concerning membership should be addressed to the Treasurer, Biological Society of Washington, National Museum of Natural History, Smithsonian Insti- tution, Washington, D.C. 20560. The Proceedings of the Biological Society of Washington is issued quarterly in February, May, August and November. Manuscripts, corrected proofs, editorial questions should be sent to the Editor, Biological Society of Washington, National Museum of Natural History, Smithsonian Institution, Wash- ington, D.C. 20560. Known office of publication: National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560. Printed for the Society by Allen Press, Inc., Lawrence, Kansas 66044 Second class postage paid at Washington, D.C., and additional mailing office. PROC. BIOL. SOC. WASH. 92(4), 1979, pp. 671-676 NOTES ON WESTERN ATLANTIC PIPEFISHES WITH DESCRIPTION OF SYNGNATHUS CARIBBAEUS N. SP. AND COSMOCAMPUS N. GEN. C. E. Dawson Abstract.—The North Atlantic is shown to be the type locality of Syng- nathus pelagicus Linnaeus. Syngnathus rousseau is a junior synonym of S. pelagicus and the so-called Caribbean Pipefish is described as S. caribbaeus n. sp. Cosmocampus n. gen. (type-species Corythoichthys albirostris Kaup) is proposed for western Atlantic species formerly referred to Corythoichthys Kaup. A review of western Atlantic pipefishes is nearing completion but some delay in publication is anticipated. Certain matters pertinent to this review have been treated previously (Dawson, 1977a, 1977b, 1978; Dawson and Allen, 1978) and this report introduces new names and information which may be of immediate interest to other workers. I here discuss the type locality and distribution of Syngnathus pelagicus, describe the presently innominate **‘Caribbean Pipefish’’ (Herald, 1965) and diagnose a new genus to accommodate western Atlantic species formerly referred to Corythoich- thys Kaup and provisionally transferred to Syngnathus Linnaeus by Daw- son (1977a). Methods follow Dawson (1977a); materials examined are de- posited in collections of the Field Museum of Natural History (FMNH), Gulf Coast Research Laboratory Museum (GCRL), Muséum National d’ Histoire Naturelle, Paris (MNHN) and National Museum of Natural His- tory, Smithsonian Institution (USNM). Syngnathus pelagicus Linnaeus Linnaeus (1758:337) described S. pelagicus from Osbeck (1757) and in- dicated the habitat, often accepted as the type locality, as “‘in Fuco na- tante.’’ Jordan and Evermann (1896) cited Osbeck’s locale as ‘‘open sea in floating seaweed,’’ whereas Weber and de Beaufort (1922), Herald (1943) and others, apparently misled by the title of Osbeck’s report, considered the type locality to be the East Indies. Reference to Osbeck (1757, 1771) shows clearly that his material was found among Sargassum (as Fucus natans or Sargazo) collected in the North Atlantic between 17.5°N, 37°21’W (of London) and 24.5°N, 39°09’W (of London) during the period 7-25 May 1752. The Sargassum Pipefish has been reported from most temperate, sub- 672 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON tropical and tropical seas but actual distribution of this species is presently uncertain. Most early records from the Indo-Pacific and southeastern At- lantic (Kaup, 1856; Duméril, 1870; Gunther, 1870) are questionable since most, if not all, specimens were probably collected from sailing ships re- turning to British or European ports. Weber and de Beaufort (1922) doubted the source of a specimen reportedly collected in the Moluccas and the origin of a specimen they report from the Celebes is also questionable. Fowler’s (1940) record from Tierra del Fuego is based on a misidentified specimen of Leptonotus blainvilleanus (Eydoux and Gervais) and I have not found Syng- nathus pelagicus among several hundred recent collections of pipefishes from New Zealand, Australia and the tropical Indo-Pacific. Studies on the distribution of this species are continuing, but present evidence suggests that S. pelagicus occurs, commonly, only in temperate-tropical Atlantic waters. Syngnathus rousseau Kaup Kaup (1856) described Syngnathus rousseau from a damaged male spec- imen sent to Paris from Martinique, without collection data, by A. Rous- seau. The unfigured description was not diagnostic, lacked information on coloration and gave counts of 16 + 34 rings, 2 + 5 subdorsal rings and 10 caudal-fin rays. Subsequently, the name has been applied to the so-called Caribbean Pipefish (Herald, 1942, 1965) which is well represented in inshore insular and mainland collections from throughout the Caribbean Sea. The holotype (MNHN 6125) is about 115 mm SL, and in poor state of preser- vation; dorsal and pectoral fins are damaged or missing and no trace of original color remains. There are 16 + 33 rings, bony preorbital is reduced to a narrow septum, the opercular ridge crosses about half of the opercle, principal body ridges are well indented between rings, and the brood pouch extends below 13 tail rings. The remaining portion of the damaged anal fin extends nearly to the rear margin of the first tail ring and, when whole, probably reached to or near the middle of the second. Despite its poor condition, the combination of 49 total rings, prominent ridges, narrow preor- bital and long anal fin indicates that the holotype of S. rousseau is conspe- cific with S. pelagicus. Since S. rousseau is a junior synonym of S. pelag- icus and there is no other available name, the following description is provided for the Caribbean Pipefish. Syngnathus caribbaeus, new species Fig. 1 Holotype.—USNM 79703 (187.0 mm SL, female), Panama, Colon, Fox Bay, 24-28 Apr. 1911, S. E. Meek and S. F. Hildebrand. Paratypes.—Panama: FMNH 8302 (1, 197); FMNH 8304 (1, 174); FMNH 673 VOLUME 92, NUMBER 4 ‘eueurg onuepYy ‘(edAjeied ‘ajewiay “TS WW 0/1) 9S61 THOD ‘Snanqqund snyjnusuks e, st LL ‘| ‘34 674 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 8305 (1, 189); FMNH 8306 (1, 203); FMNH 8307 (1, 180); GCRL 1956 (1, 170); GCRL 14541 (1, 141); USNM 79697 (1, 191); USNM 79701 (2, 186— 186.5). Venezuela: GCRL 15524 (2, 53-163); GCRL 15525 (1, ca. 190). Diagnosis.—Total rings 48-52; total subdorsal rings 5.5—7.25; snout length averages 1.9 in head length; bony preorbital typically moderate, nei- ther broad nor reduced to a narrow septum; anal fin short, reaches little beyond anterior margin of 2nd tail ring; without prominent dark bands on body. Description.—Rings 15-18 + 31-35 (usually 17 + 33), dorsal-fin rays 25— 32 (usually 28-30), subdorsal rings 2.5-1.0 + 3.5—5.75 (usually 2.0-1.5 + 4.25—5.0), pectoral-fin rays usually 13-14, caudal-fin rays 10, anal-fin rays 3. Counts and measurements (mm) of 187.0 mm SL, female, holotype (USNM 79703) follow: rings 17 + 34, dorsal-fin rays 29, subdorsal rings 1.5 + 5, pectoral-fin rays 14 (2), head length 23.0, snout length 12.0, snout depth 2.2., length of dorsal-fin base 19.3, anal ring depth 6.3, trunk depth 7.2, pectoral-fin length 4.3, length of pectoral-fin base 3.9. In alcohol, the holo- type is mainly brown and there are traces of brownish bands on the dorsal fin. Other material often with brown lateral stripe on snout; body plain, mottled or with faint pale bars; dorsal fin plain or with indistinct brown bands in both sexes. Comparisons.—Among western Atlantic congeners, this species (as noted by Herald, 1942, 1965) is similar to S. floridae and S. pelagicus in most meristic and morphometric features. It is best separated from S. floridae by the moderate width of bony preorbital (typically broad in floridae) and by a lower pectoral-fin length in HL ratio (averages 6.2 against 7.8). The mod- erate preorbital width and shorter anal fin separate S. caribbaeus from S. pelagicus wherein the preorbital is narrow or septum-like and the anal fin usually reaches to or beyond the middle of the 2nd tail ring. In addition, adult females have a slender trunk (deep in pelagicus), brooding males are infrequent under 100 mm SL (common at 75-80 mm in pelagicus) and S. caribbaeus reaches a larger size (at least 225 mm SL against ca. 180 in pelagicus). This species commonly frequents shallow inshore waters where- as S. pelagicus (except following storms) usually occurs in offshore or open sea collections. Etymology.—Named caribbaeus, an adjective referring to the Caribbean distribution of the species. Distribution.—Known from the Greater and Lesser Antilles and mainland coasts of Central and S. America from Belize to Venezuela. Cosmocampus, new genus Type-species.—Corythoichthys albirostris Kaup 1856. Diagnosis.—Superior trunk and tail ridges discontinuous near rear of dor- VOLUME 92, NUMBER 4 675 sal fin, inferior trunk and tail ridges continuous, lateral trunk and tail ridges discontinuous below dorsal-fin base. Snout length 1.7-3.8 in HL; median dorsal snout ridge low, entire to denticulate; median lateral snout ridge present or absent; median dorsal head ridges distinct to strongly elevated; supraopercular ridge present; opercle with complete or incomplete median ridge, usually prominent, often angled dorsad and margined with radiating striae; pectoral-fin base usually with 2 prominent ridges. Principal body ridges prominent, occasionally strongly elevated; ridge margins indented to deeply notched between rings; juvenile and adults often with posterior an- gles of tail rings produced as short spines; dermal flaps typically present in juveniles or adults. Trunk rings 15-18, total rings 40-57, dorsal-fin rays 19— 27, pectoral-fin rays 10-15, anal-fin rays 2—4, caudal-fin rays typically 10. Dorsal-fin origin usually on trunk, fin base not elevated, total subdorsal rings 4.25—6.25. Brood pouch below 12-20 tail rings; pouch plates present; pouch closure not the inverted type of Herald (1959); without odontoid processes (Dawson and Fritzsche, 1975). Comparisons.—The principal body ridge configuration of Cosmocampus is shared with several syngnathine (tail pouch) genera. This genus differs from Syngnathus sensu stricto in the presence of a supraopercular ridge and dermal flaps and in the absence of an inverted pouch closure. Cosmocampus differs from Bryx Herald in possessing an anal fin, from Corythoichthys Kaup in the presence of pouch plates and from Bhanotia Hora in lacking bony inclusions in opercular membranes and inverted pouch closure. Fi- nally, Cosmocampus is differentiated from two nominal Australian genera by the absence of the elevated plate-like snout ridge characteristic of His- tiogamphelus McCulloch, and by moderate snout length and lower trunk ring count (15-18 against 25—26) from the long-snouted monotypic Hypse- lognathus Whitley. Etymology.—From the Greek kosmos (ornament or decoration) and kam- pos (sea-animal), in allusion to the ridges and dermal flaps decorating the head of the type-species; gender, masculine. Remarks.—TYwo other western Atlantic species, Syngnathus brachy- cephalus Poey 1868 and Corythoichthys profundus Herald 1965, are also re- ferred to Cosmocampus. Acknowledgments I thank M. L. Bauchot (MNHN) for access to material in her care and A. C. Wheeler [British Museum (Natural History)] for sharing his knowledge of Linnaean specimens. This study was supported in part by National Sci- ence Foundation Grant BMS 75-19502. 676 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Literature Cited Dawson, C. E. 1977a. Review of the pipefish genus Corythoichthys with description of three new species.—Copeia 1977:295-338. . 1977b. Synopsis of syngnathine pipefishes usually referred to the genus Ichthyocam- pus Kaup, with description of new genera and species.—Bull. Mar. Sci. 27:595—650. . 1978. Micrognathus vittatus (Kaup), a junior synonym of M. crinitus (Jenyns), with description of the insular pipefish, M. tectus new sp.—Copeia 1978:13-16. , and G. R. Allen. 1978. Synopsis of the ‘‘finless’’ pipefish genera (Penetopteryx, Ap- terygocampus and Enchelyocampus, gen. nov.).—Rec. West Austr. Mus. 6:391—411. , and R. A. Fritzsche. 1975. Odontoid processes in pipefish jaws.—Nature 257:390. Dumeril, A. 1870. Histoire naturelle des poissons ou ichthyologie générale. Tome second.— Ganoides, dipnés, lophobranches. Paris. 624 pp. Fowler, H. W. 1940. The fishes obtained by the Wilkes Expedition, 1838—1842.—Proc. Amer. Phil. Soc. 82:733-800. Giinther, A. 1870. Catalogue of the fishes of the British Museum.—Taylor and Francis, Lon- don. Vol. 8. 549 pp. Herald, E. S. 1942. Three new pipefishes from the Atlantic coast of North and South America, with a key to the Atlantic American species.—Stanford Ichthyol. Bull. 2:125—-134. 1943. Studies on the classification and relationships of the American pipefishes.— Ph.D. Dissertation, Stanford Univ. 339 pp. . 1959. From pipefish to seahorse—a study of phylogenetic relationships.—Proc. Calif. Acad. Sci., 4th Ser., 29:465—473. . 1965. Studies on the Atlantic American pipefishes with descriptions of new species.— Proc. Calif. Acad. Sci., 4th Ser., 32:363-375. Jordan, D. S., and B. W. Evermann. 1896. The fishes of North and Middle America.—Bull. U.S. Nat. Mus. 47. Kaup, J. 1856. Catalogue of lophobranchiate fish in the collection of the British Museum.— Taylor and Francis, London. Linnaeus, C. 1758. Systema Naturae etc., 10th ed., Vol. 1, 824 pp. Osbeck, P. 1757. Dagbok 6fver en Ostindisk resa etc.—L. Grefing, Stockholm. . 1771. A voyage to China and the East Indies (English transl. of 1757 report by J. R. Forster). Vol. 2.—Benjamin White, London. Weber, M., and L. F. de Beaufort. 1922. The fishes of the Indo-Australian Archipelago.—E. J. Brill, Leiden. 4:1-140. Gulf Coast Research Laboratory Museum, Ocean Springs, Mississippi 39564. PROC. BIOL. SOC. WASH. 92(4), 1979, pp. 677-681 ON THE TYPES IN THE GENUS PELOSCOLEX LEIDY (OLIGOCHAETA: TUBIFICIDAE) Ralph O. Brinkhurst Abstract.—The selection of neotypes for Peloscolex variegatus is de- fended, but the subdivision of the genus is supported. Four groups of species are emerging, but the final disposition of all former Peloscolex species cannot be ascertained as yet. When the author first undertook to revise the Tubificidae it rapidly be- came apparent that few genera, if any, were based on properly defined type species. Several names had become fixed in the literature by common usage, and so a deliberate attempt was made to preserve those names so long as this action did not clearly violate established nomenclatural procedures. The genus Peloscolex was among the most difficult to deal with as, unlike most other genera in the family, it seemed to be characterized by the coating of papillae on the body wall which led authors to assume that many species were congeneric without detailed description of the male reproductive struc- tures or even the setae where these were obscured by the papillae. The type species is P. variegatus Leidy, 1851, by reason of monotypy, but it has never been recorded since it was first discovered. It would have been possible to regard Leidy’s description as inadequate, and to have removed the genus name from availability, but this would almost certainly have been challenged by some subsequent author claiming to be able to recognise the type from the brief original account, which was limited to the following details: — Setae usually 10 in each anterior dorsal bundle, sometimes 6. Ventral setae bifid. — Prominent or elevated rounded tubercles in transverse circles. — From a ferruginous spring, Philadelphia. At an early stage in the investigation of North American Tubificidae the author came across some hitherto undescribed material which seemed rea- sonably consistent with the original account of variegatus, and selected a neotype (USNM 32626) and paraneotypes (USNM 32627, American Mu- seum of Natural History 3662, British Museum Natural History 1964.15.8) as noted in Brinkhurst (1965). This designation was deficient in that it re- ferred back to the redescription by Brinkhurst (1962) but did not mention that a search of the collections at Philadelphia proved the types had been 678 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON destroyed. This action preserved the name Peloscolex, although it has be- come generally acknowledged that the papillate body wall is no longer a good generic character and that some division of the genus is required. Holmquist (1978) has challenged the selection of the neotypes, which could certainly be shown to fail a stringent application of Article 75 of the Inter- national Code of Zoological Nomenclature, and claims to be able to rec- ognize another American species, P. multisetosus (Smith, 1900), as con- specific with Leidy’s original. The evidence for this will be examined point by point, and the implications of the decision will be discussed before some suggestions are made as to the necessary division of the species assemblage into genera. The setae.—Leidy’s specimen(s) had 6-10, usually 10 setae per bundle dorsally, bifid setae ventrally. Brinkhurst’s specimens had 2—4 hair setae and 2—4 short hair-like setae in dorsal bundles for a total of 4-8; the ventral setae are bifid apart from those of II-IV where at least one may be simple-pointed, though this is very difficult to see without specifically looking. Holmquist reports (1978) that Brinkhurst’s types have ‘‘only up to about 3 long hair setae and one or a few short pectinate setae in the dorsal bun- dles,’’ and she suggests that Leidy ‘‘would hardly have seen more than double the number of long dorsal hair setae... .”’ The description of P. multisetosus lists 3-14 hair setae and 1-5 lyre- shaped pectinate setae for a total of up to 19 setae per bundle, or about double the number Leidy saw! As Leidy did not mention any differences between the dorsal setae, I have assumed the hairs and hair-like setae were not separated by him. As he could see the ventral bifids he most certainly would have seen the obvious pectinate setae in what was later described as multisetosus. The papillae.—The early literature on Peloscolex talks about two types of papillae, body wall extensions through the foreign matter called sensory or secretory papillae, and the general papillae with their included foreign matter. Holmquist (1978) illustrates these well. The sensory papillae are hard to see in preserved worms, in which they may be expected to be affected by rough preservation methods often employed. Hrabe (1973) dis- cussed these in P. velutinus (Grube), a species clearly close to variegatus sensu Brinkhurst as noted by Holmquist (1978) who suggests grouping these and other taxa in the genus Embolocephalus Randolph. In velutinus there are, according to Hrabe (op. cit.) one line of large secretory papillae on the anterior segments of the transverse line of the setal bundles, and posteriorly two lines with one between the setal lines. He goes on to state that the younger worms have secretory papillae of only one size, so I take it that mature forms have two sizes of these without considering the ordinary pa- pillae (which are smaller than the large secretory ones). Kasprzak (1976) VOLUME 92, NUMBER 4 679 published a photograph of the papillae of P. velutinus plus what he refers to as cutaneous glands. The papillae are in 2—3 regular transverse rows, and are prominent. If these specimens are conspecific with my velutinus mate- rial, their appearance is relevant to this argument. In my collection of few specimens of either velutinus or my variegatus I cannot see these elevated (enlarged) papillae on the whole mounts. It is my contention that the de- scriptions of papillae by Hrabe and Kasprzak are almost identical to that of Leidy, and that other similarities between velutinus and variegatus validate the comparison. The discovery by Holmquist of spermathecal setae in var- iegatus requires a close comparison of this with velutinus and _ nikolskyi Last. and Sok. The papillae in multisetosus are of a very different dimension. They are so remarkable that Leidy may well have been moved to describe them in much stronger terms than prominent or elevated. Factually, both multise- tosus and variegatus mihi have papillae which qualify, they differ in size and ease of visibility of the sensory/secretory papillae, but variegatus usu- ally has a closer covering of the body wall papillae not specifically men- tioned by Leidy. Locality.—Holmquist does not discuss locality or habitat although these points were raised in correspondence with her. Philadelphia is within the geographical range of both variegatus mihi and multisetosus (J. Hiltunen, personal communication). The species overlap in the St. Lawrence Great Lakes but multisetosus seems to be a more southern species at its northern limits with the reverse for variegatus. Specimens resembling this species have been found in Washington State, most recently in Mowitch Lake, Mount Rainier Park, 1966 by G. Larson. These specimens are notable in that there are up to 5 (serrate) hair setae and 6 short hair-like setae with minutely pectinate tips, but there are usually only 3-4 of each. In segment II the dorsal bundle includes a less hair-like bifid seta with an elongate upper tooth. These also resemble P. nikolskyi, known from Asiatic Russia, and all of these may be conspecific. Habitat.—A ferruginous spring suggests to me a close connection to ground water and adequate aeration. P. multisetosus is often found in warm, eutrophic habitats if not frankly polluted sites. P. variegatus, on the other hand, is acold water oligotrophic species largely limited to big lakes. Similar ecological habits in species like L. profundicola and R. falciformis have led to their detection in large lakes and small spring-fed streams, so a spring as the type locality seems consistent with the known ecology of variegatus mihi. Mr. M. Loden (personal communication) collected variegatus in cold Pennsylvania streams in winter in sand-gravel substrates. Taxonomic stability.—Several possible decisions could flow from the var- ious positions adopted. Leidy’s species could be declared to be unrecog- nizable by virtue of the impossibility of deciding between multisetosus 680 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Smith and variegatus mihi on the basis of Leidy’s description which was, to say the least, inadequate. If Leidy’s species were to be dubbed a species inquirenda I assume the generic name would be unavailable if it were placed on the Official Index by the Commission. The whole lexicon of old generic names could then be reactivated including Saenuris Grube, Embolocephalus Randolph, Spirosperma Eisen, Pachydermon Claparede, Hemitubifex Bed- dard and TJubificoides Lastockin (see Brinkhurst and Jamieson 1971 for details). Indeed, this is exactly what Holmquist is obliged to do, although Saenuris, Pachydermon, Peloryctes and Hemitubifex for Edukemius are not available as can be deduced by inspection of the synonymies in Brink- hurst and Jamieson, 1971, pp. 453, 481, 506, 508-9, 572-74. As the genus is to be broken up, rejection of Peloscolex might seem to be the wisest course, if legal, but stability could be challenged at any time by a claimed recognition of Leidy’s species. By accepting the author’s decision, the name Peloscolex remains avail- able for the major group of species formerly in that assemblage. Accepting P. multisetosus as a synonym of variegatus would cause max- imum confusion in a literature now very extensively used by applied biol- ogists using sludge-worms in pollution biology in North America. The grounds for doing this would have to be very substantial for it to gain ac- ceptance by this community, which is continually disturbed by changes in generic names. Holmquist suggests that Smith (1900) suggested a possible synonymy of his multisetosus and P. variegatus of Leidy, but my interpre- tation of that suggestion is that Smith believed the two to be potentially congeneric though he described his species as an Embolocephalus. Beddard (1895) stated categorically that variegatus of Leidy had dorsal setae all hair- like, as in velutinus, but it is not clear that he saw types or communicated with Leidy. The latter was President of the Philadelphia Academy of Na- tional Sciences (where his collections were) until he died in 1891. Discussion.—Holmquist has contributed two major changes to the de- scription of variegatus mihi. She discovered genital setae of a form that I anticipated might be present when discussing the closely related nikolskyi Last. and Sok. (Brinkhurst and Jamieson 1971, p. 508), but I hardly expected my prophesy to be substantiated from my own neotypes. She also finds no cuticular penis sheaths which I believed to be present. As these are absent in the related velutinus and nikolskyi it would seem that I was looking at the normal cuticular layer of the penis sac, which becomes thicker in gen- uine penis sheaths. Otherwise I suggest that my variegatus is no more at odds with Leidy’s original than is multisetosus. My suggestion to Holmquist in a personal communication was that a sensible option might be to declare Leidy’s de- scription to be inadequate and to reject the name Peloscolex, especially as the genus is bound to be divided as a result of more recent work by a number of authors. If my variegatus is upheld, the name Peloscolex can be VOLUME 92, NUMBER 4 681 used for the bulk of the limnic species (which Holmquist divides into Em- bolocephalus, Spirosperma, Orientodrilus and perhaps more—she does not yet attribute all the species she has seen into genera, nor does she mention at least six species in the recent literature). The name Embolocephalus Randolph is not available for multisetosus and its ally moszynskii as the type of the genus is velutinus Grube, so that a new name must be created. If Holmquist’s decision is adopted, two well known North American species swap names (variegatus for multisetosus), variegatus gets a new name as yet unspecified, and the generic name Peloscolex becomes applied to a pair of species that are clearly separate from the genus as currently defined. I can only conclude that Holmquist has chosen the worst possible path from the point of view of stability of the nomenclature, and I would prefer to retain my designation of neotypes of variegatus. The other decisions made by Holmquist are as yet incomplete, but it can be said that the two genera Tubificoides and Edukemius sensu Holmquist are part of a larger assemblage of marine species (including former Tubifex species and new entities) currently being assembled by Brinkhurst and Bak- er (1979) the type species of which should correctly be cited as T. het- erochaetus Lastockin, 1937 (synonym = T. swirencowi Jaroschenko, 1948 = Peloscolex swirencowi Jar. Hrabe, 1964; non Limnodrilus heterochaetus Michaelsen, 1926 = Peloscolex heterochaetus [Mich.] de Vos 1936;—all vide Brinkhurst and Jamieson 1971, pp. 508, 511, 521 for references). Literature Cited Beddard, F. E. 1895. Monograph of the order Oligochaeta.—Clarendon Press, Oxford. Brinkhurst, R. O. 1962. A re-description of Peloscolex variegatus Leidy (Oligochaeta, Tub- ificidae) with a consideration of the diagnosis of the genus Peloscolex.—Int. Revue Gesamten Hydrobiol. 47:301-—306. 1965. Studies on the North American Aquatic Oligochaeta II. Tubificidae.—Proc. Acad. Nat. Sci. Philad. 117:117-172. , and H. R. Baker. 1979. A Review of the Marine Tubificidae (Oligochaeta) of North America.—Can. J. Zool. 57:1553—1569. , and B. G. M. Jamieson. 1971. Aquatic Oligochaeta of the World.—Oliver and Boyd, Edinburgh. Holmquist, C. 1978. Revision of the Genus Peloscolex.—Zool. Scripta 7: 187-208. Hrabe, S. 1973. Ona collection of Oligochaeta from various parts of Yugoslavia.—Biol. Vest. Lubl. 21:39-S0. Kasprzak, K. 1976. Contribution to the knowledge of aquatic Oligochaeta of Italy.—Acta Zool. Cracoviensia 21:331-350. Leidy, J. 1851. Description of new genera of Vermes.—Proc. Acad. Nat. Sci. Philad. 1850- 51:5:124—-126. Smith, F. 1900. Notes on species of North American Oligochaeta. III. List of species found in Illinois and description of Illinois Tubificidae.—Bull. Ill. St. Lab. Nat. Hist. 5:441- 458. Institute of Ocean Sciences, P.O. Box 6000, Sidney, British Columbia, Canada V8L 4B2. PROC. BIOL. SOC. WASH. 92(4), 1979, pp. 682-688 A LARGE PARASUCHIAN (PHYTOSAUR) FROM THE UPPER TRIASSIC PORTION OF THE CULPEPER BASIN OF VIRGINIA (USA) Robert E. Weems Abstract.—Postcranial remains of a large parasuchian comparable to Ru- tiodon manhattanensis were recovered in an excavation at Dulles Interna- tional Airport, Fairfax County, Virginia. While fragmentary, these remains represent the first parasuchian known from the Culpeper Basin and consti- tute the only tetrapod skeletal material reported from this basin; previous reptile records are based on footprints. The problems of taxonomic assign- ment of the known Newark parasuchian remains is briefly discussed. Although the Culpeper Basin is one of the larger of the exposed Newark basins comprising the Newark Supergroup (Olsen, 1978), until recently it received relatively little attention except for the pioneering mapping done by Roberts (1928). In the last few years a number of papers have appeared on local problems within the basin but only a few have expanded on the lithostratigraphy and chronostratigraphy put forth by Roberts. Cornet et al. (1973) and Cornet (1977) established the existence of Jurassic (Liassic) age sediments toward the top of the basin’s sequence; prior to this discovery the entire column was presumed to belong within the upper Triassic. These conclusions have been reinforced by similar interpretations of the Culpeper basin fish fauna by Schaeffer and McDonald (1978). Major changes in the basin lithostratigraphy have been proposed by Lee (1977), who has proposed a series of names for units in the basin quite different from those used (in part erroneously) by Roberts. The remains described herein were excavated by D. H. Dunkle and R. E. Eggleton in 1959 from a drainage ditch due east of the south end of the main north-south runway at Dulles International Airport (Figure 1). On the presumption that about 30,000 stratigraphic feet of sediment are present in the basin, this locality lies about 5,000 stratigraphic feet (1,600 meters) above the base of the basin’s column and about 15,000 stratigraphic feet (4,700 meters) below the lowest lava flow in the basin (which falls slightly above the Triassic-Jurassic systemic boundary [Cornet, 1977]). This places the specimen near the base of the Bull Run Shale as defined by Roberts and near the base of the Balls Bluff Siltstone as defined by Lee. The Dulles specimen (USNM 22381) was found in a red calcareous mud- VOLUME 92, NUMBER 4 683 International Airport / Fig. 1. Map showing location of Dulles International Airport and the area within it from which the parasuchian skeleton was recovered (black diamond). Horizontal rule—pre-Tniassic metamorphic terrane; vertical rule—sediments of known Jurassic age lying above basal lava flow. Since basal flow is slightly above the Triassic-Jurassic boundary and the exact position of this boundary is not yet established, most but not all of the unruled area is of late Triassic age. Stippled region on Virginia base map is the approximate area underlain by the Culpeper basin. 684 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON stone with large white calcareous lumps probably representing caliche con- centrations or possibly oncalites. The bones are nearly white and contrast sharply with the surrounding matrix. The remains include a posterior cer- vical vertebra, a fragmentary dorsal vertebra, vertebral fragments, rib frag- ments and polygonal armor plates. These clearly pertain to a very large parasuchian*. Even though the Dulles specimen is extremely fragmentary, because it is the first evidence for parasuchians reported from the Culpeper basin (and indeed the only tetrapod skeletal remains of any kind reported from the basin) it is still an important find. Although this specimen was passingly mentioned by Eggleton (1975), based on its identification as a parasuchian by Nicholas Hotton III of the Smithsonian Institution, so far no detailed, widely available account has been published. Since parasuchi- ans are universally accepted as good indicators of Late Triassic age, this specimen offers independent evidence to support the contention of Cornet (1977) that much of the lower Culpeper basin belongs within the upper Triassic system as has been traditionally assumed. Gregory (1962a) undertook a major revision of the parasuchia which re- sulted in the strong reduction of a plethora of generic names to only four applicable to American material: Angistorhinus, Paleorhinus, Rutiodon, and Phytosaurus. Among the species which he accepted as valid, only a few assignable to Rutiodon and Phytosaurus equal the size of the Dulles specimen. Since the irregularly polygonal and subequant shapes of the ar- mor plates preclude placement of the Dulles specimen in the genus Phyto- saurus as currently defined (Baird, as acknowledged in Colbert, 1965) and since all parasuchian material so far known from elsewhere in the Newark Supergroup seems referable to Rutiodon (Colbert, 1965) the Dulles speci- men can be placed within this genus with some confidence. At the species level, the status of Newark parasuchians is thoroughly unclear. Lea (1851) was the first to describe a fragmentary, large parasu- chian skeleton from the Pennsylvania Gettysburg basin, which he named Clepsysaurus pennsylvanicus. Soon after, Emmons described two smaller species, Rutiodon carolinensis (1856) and Rutiodon sulcatus (1857), on the basis of fragmentary material from the North Carolina Deep River basin. Both Rutiodon types are now in the United States National Museum col- lections. The type of R. sulcatus is distinctive; it is a valid taxon and an archosaur, but almost certainly it is not a parasuchian. Emmons (1860) later described more complete cranial remains of R. carolinensis and McGregor (1906) described most of the postcranial skeleton based on bones from nu- merous individuals recovered in the type area. As a result of these papers, as well as Gregory’s significant re-analysis of the skull (1962b), R. caroli- * Parasuchians have commonly also been called phytosaurs, but the latter name recently has been considered less proper (Chatterjee, 1978). VOLUME 92, NUMBER 4 685 CMS. Fig. 2. A, Lateral view of best preserved cervical vertebra from the Dulles parasuchian, slightly distorted from compression; B, Armor plate recovered with the Dulles parasuchian. Maximum length is 10.5 cm.; C, Armor plate recovered with the Dulles parasuchian. Maximum length, 6.3 cm. nensis 1S now fairly well known, assuming all parts are correctly assigned to only one species. A second skull from the Deep River basin, described by Marsh (1896) as *“Rhytidodon rostratus,’’ is almost certainly also refer- able to R. carolinensis (Gregory, 1962b). In 1913, Huene described remains of another, much larger parasuchian from the Palisades bluff on the Hudson River near New York City as Rutiodon manhattanensis. A few years later Sinclair (1918) described fragments of yet another very large parasuchian from Pennsylvania which he chose to refer to Huene’s R. manhattanensis. Neither specimen included a skull, though a fair portion of the postcranial skeleton was represented. Since the best and most reliable diagnostic char- acters are in the skull (Gregory, 1962a), at present only large size really separates R. manhattanensis from R. carolinensis. Thus Colbert, when he described a small skull from the Lockatong Formation of New Jersey at Fort Lee (1965) was compelled to assign it to Rutiodon carolinensis (with which it is fully comparable) rather than to a young R. manhattanensis (for which the skull is unknown), while acknowledging the impossibility of mak- ing too firm a decision in view of how poorly R. manhattanensis is known. Camp (1930) and Colbert and Chaffee (1941) accepted Clepsysaurus penn- 686 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 3. (Left) Posterior view of the cervical figured in Fig. 2A; (Right) anterior view of the cervical figured in Fig. 2A. sylvanicus as a valid species, but more recently the trend has been to syn- onymize it with R. manhattanensis which constitutes a much more diag- nostic type (see Colbert [1965] for a thorough discussion of this problem). Thus Clepsysaurus pennsylvanicus has been relegated to the status of a nomen dubium. Yet even the validity of Rutiodon manhattanensis is none too certain. Only size presently distinguishes R. manhattanensis from R. carolinensis. It is entirely possible that some environmental bias in the New- ark basins causes us to find only young or only old specimens of Rutiodon but not both together; perhaps Rutiodon was a swamp or lake dweller as a juvenile and an upland dweller as an adult (an idea partially developed by Gregory, 1962a, p. 688). If so, R. manhattanenesis and R. carolinensis may really be only different age groups of the same species. Thus either one or two species of Rutiodon are known to be present in the Newark Supergroup, depending upon the preferences and biases of the worker. In view of the existence of this state of taxonomic ambiguity, no purpose would be served by firmly assigning the Dulles material to either existing species or to a new species. Though stratigraphically significant, it is best for now to assign this specimen to Rutiodon cf. manhattanensis until new and much more com- plete material from a number of the Newark basins permits a fresh and meaningful look at the intricacies of Newark parasuchian taxonomy. VOLUME 92, NUMBER 4 687 Acknowledgments The author wishes to gratefully acknowledge the following individuals for kindly reading and criticizing the manuscript for this paper: Dr. Donald Baird (Princeton University), Dr. Nicholas Hotton III (U.S. National Mu- seum), Dr. K. Y. Lee (U.S. Geological Survey) and Dr. Roy C. Lindholm (George Washington University). Literature Cited Camp, C. 1930. A study of the phytosaurs with descriptions of new material from North America.—Mem. Univ. California 10:1—174. Chatterjee, S. 1978. A primitive parasuchid (phytosaur) reptile from the Upper Triassic Maleri Formation of India.—Palaeontol. 21(1):83-127. Colbert, E. 1965. A phytosaur from North Bergen, New Jersey.—American Mus. Novitates 2230:1-25. , and Chaffee, R. 1941. The type of Clepsysaurus pennsylvanicus and its bearing upon the genus Rutiodon.—Notulae Nat., Acad. Nat. Sci. Philadelphia 90:1-19. Cornet, B. 1977. The palynostratigraphy and age of the Newark Supergroup.—Ph.D. disser- tation, the Pennsylvania State Univ., Univ. Park, Pennsylvania, 505 p. , Traverse, A., and McDonaid, N. 1973. Fossil spores, pollen and fishes from Con- necticut indicate Early Jurassic age for part of the Newark Group.—Science 182:1243- 1247. Eggleton, R. E. 1975. Preliminary geologic map of the Herndon quadrangle, Virginia.—U.S. Geol. Survey open-file report 75-386, scale 1:24,000. Emmons, E. 1856. Geological Report on the Midland Counties of North Carolina.—G. P. Putnam and Co., Raleigh, N.C., 352 p. . 1857. American Geology, containing a Statement of the Principles of the Science, with full Illustrations of the Characteristic American Fossils, with an Atlas and a Geo- logical Map of the United States, Part 6.—Albany, N.Y., 152 p. . 1860. Manual of Geology, 2nd ed.—New York. Gregory, J. 1962a. The genera of phytosaurs.—American Jour. Sci. 260:652-690. . 1962b. The relationships of the American phytosaur Rutiodon.—American Mus. Nov- itates 2095:1-22. Huene, F. von. 1913. A new Phytosaur from the Palisades near New York.—Bull. American Mus. Nat. Hist. 32(15):275-284. Lea, I. 1851. Remarks on Clepsysaurus pennsylvanicus.—Proc. Acad. Nat. Sci. Philadelphia 5:205. Lee, K. Y. 1977. Triassic stratigraphy in the northern part of the Culpeper basin, Virginia and Maryland.—United States Geol. Survey Bull. 1422-C, 17 p. Marsh, O. 1896. A new belodont reptile (Stegomus) from the Connecticut River sandstone.— American Jour. Sci., ser. 4, 2:59-62. McGregor, J. 1906. The phytosauria, with especial reference to Mystriosuchus and Rutio- don.—Mem. American Mus. Nat. Hist. 9:29-101. Olsen, P. 1978. On the use of the term Newark for Triassic and Early Jurassic rocks of eastern North America.—Newsl. Stratigr. 7(2):90-95. Roberts, J. 1928. The geology of the Virginia Triassic.—Virginia Geol. Survey Bull. 29: 1-205. Schaeffer, B., and McDonald, N. G. 1978. Redfieldiid fishes from the Triassic-Liassic Newark Supergroup of Eastern North America.—Bull. American Mus. Nat. Hist. 159(94):129- 174. 688 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Sinclair, W. 1918. A large parasuchian from the Triassic of Pennsylvania.—American Jour. Sci., ser. 4, 45:457—462. Stop 928, U.S. Geological Survey, Reston, VA 22092. PROC. BIOL. SOC. WASH. 92(4), 1979, pp. 689-696 COMMENTS ON THE PHYLOGENY OF PERCHING BIRDS Alan Feduccia Abstract.—The bony stapes (columella) is a unique character in birds in that the primitive condition is the retained reptilian condition of the element, and derived ‘‘pockets’’ of stapedial morphologies occur that may be of importance in clarifying certain phylogenetic relationships. Scanning elec- tron micrographs (SEM’s) of various views of stapes of suboscines, certain coraciiforms and trogons are here presented and reveal with clarity the manifest differences betweeen suboscines and the alcediniform coraci- iforms. These new data, combined with new data from sperm morphology of suboscines, suggests that the Passeriformes is a monophyletic assem- blage, that the suboscine and alcediniform stapes evolved independently, and that the suboscines are more closely related to the oscines than to the alcediniforms. In a previous paper (Feduccia, 1977), I proposed a hypothetical phylogeny of the passerines and related avian groups, birds that have presented difficult phylogenetic problems because of their morphological uniformity and the probability of massive convergent evolution. My reconstruction was based primarily on the discovery of a new avian taxonomic character, the config- uration of the bony stapes or columella, which had until recently been over- looked because of its minute size (one to several mm) and its location in the recesses of the middle ear cavity. In addition, it is often broken or lost in skeletal preparations. The avian stapes is perhaps unique as an avian morphological character in that the primitive condition is the retained rep- tilian stapedial morphology, which is characteristic of most groups of birds. In some groups, however, the stapes exhibits peculiar derived morpholo- gies. Where these ‘‘pockets’’ of derived stapedial morphologies are found they are considered as strong indications of evolutionary affinity, unless there are compelling reasons to assume that the morphological similarities are due to convergent evolution. I made the following broad conclusions (Feduccia, 1977). First, the hoo- poes (Upupidae) and wood-hoopoes (Phoeniculidae) are monophyletic with- in the Coraciiformes (sensu Wetmore 1960); previously there was consid- erable dispute as to their relatedness. These two families are characterized by a stapedial morphology that is termed the ‘‘anvil’’ stapes and is illustrated in Fig. 1. Second, the coraciiform families containing the bee-eaters (Mer- 690 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 1. Views along the lengths of the bony stapes of a hoopoe (Upupa epops), left; and a wood-hoopoe (Phoeniculus purpureus), right. SEM’s x 40, and reduced here x 1/3. opidae), kingfishers (Alcedinidae), motmots (Momotidae), and todies (To- didae) all possess a peculiar derived stapedial morphology characterized by a bulbous footplate (with particular conformations; see Feduccia, 1975) that is shared with the trogons (Trogonidae). I concluded that this bee-eater/ kingfisher/motmot/tody assemblage is monophyletic, and that the trogons are allied with it. Trogons had until then been placed in a monotypic order, the Trogoniformes, and there was little but speculation as to their phylo- genetic relationships. I proposed that these families (including the trogons) be combined in a separate order Alcediniformes close to the old order Cor- aciiformes, clearly their sister group. The separation of the Alcediniformes as a distinctive order is a matter of personal taxonomic preference. Figure 2 illustrates three views of the stapes of a kingfisher, Ceryle rudis, and a trogon, Priotelus temnurus. There is more stapedial variation within the families of the coraciiform ‘‘alcediniforms’’ than between the trogon and kingfisher illustrated here. The third conclusion, which now demands modification in view of new evidence that has recently come to light, was that the suboscines (classically the suborder Tyranni of the Passeriformes) and the oscines (Passeres) did not share an immediate common ancestor, and that the classical order Pas- seriformes was not monophyletic. The point focal to the discussion of passerine relationships involves the fact that while the morphologically uniform (and presumably more advanced structurally) oscines retain the primitive condition of the stapes, the subos- cines have a derived stapedial morphology characterized by a bulbous foot- plate region with certain peculiarities. Suboscines, structurally more prim- VOLUME 92, NUMBER 4 691 itive than oscines, are a real ‘‘South American’’ group that has been eliminated in the Old World (presumably by competition with oscines) ex- cept for two highly specialized, small families, the Pittidae and Eurylaimi- dae, and some relicts restricted to Madagascar. My point was that although the suboscines and oscines exhibit many osteological and other similarities, the stapes of suboscines was more similar to that of the ‘‘alcediniforms”’ than to the oscines (though different from the former). Since then, however, information has forced me to view the hypothesis of a polyphyletic order Passeriformes as improbable. Scanning electron micrographs (SEM’s) of suboscine and alcediniform stapes reveal many differences, especially in the footplate region (Figs. 2 and 3), which do not argue for homology of the two and, in fact, would seem to indicate a high probability that the two morphologies evolved in- dependently. In the alcediniforms there is a relatively much larger footplate fossa that is more rounded and bulbous in appearance. The major difference, however, is seen in the conformation of the footplate (Fig. 3). In the alce- diniforms the region of the footplate near the shaft of the stapes is smoothly rounded, so that when these specimens are placed in a depression slide filled with liquid, the Jarge fossa will face upward. In the suboscines the same region tapers to a point, so that when suboscine stapes are placed in a similar preparation the fossa turns and lies to one side. We (Henley et al., 1978) have revealed characters in oscine sperm that make the group perfectly definable. Oscines have non-motile sperm that occur in bundles in the testes when placed in warm saline; other birds have non-bundled motile sperm within the testes under similar conditions. In addition, in oscine sperm cross sections are characterized by a tripartite structure of an undulating membrane. The components are a relatively straight axoneme with the 9 + 2 arrangement of the microtubules, a helically wound strand of mitochondria, and a longitudinal array of singlet microtu- bules. These sperm are highly derived and unique among vertebrates. At the time of our study we had not examined suboscine sperm, but I have recently discovered that suboscines have the sperm bundles (Fig. 4) char- acteristic of, and previously thought to be unique to, the oscines. Only two species, the Eastern Wood Pewee (Contopus virens) and the Eastern King- bird (Tyrannus tyrannus) have been examined, but because of the uniformity observed and the extreme difficulty in making these preparations, it was felt that the data should be presented here. The ultrastructure of suboscine sperm (Fig. 5) while showing an undulat- ing membrane, is different in many important details from that of oscines. Suboscine sperm differ in the geometry of the microtubules that surround the axoneme. The axoneme is separated from the bundle of singlet micro- tubules by double plasma membrane as indicated in Fig. 5, and the entire unit is surrounded by another plasma membrane. In addition, the singlet PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 2. Three views along the lengths of the bony stapes of: right, a trogon (Trogonidae: Priotelus temnurus), middle, a kingfisher (Alcedinidae: Ceryle rudis), and left, a suboscine (Cotingidae: Rupicola peruviana). SEM’s were taken so as to have all to the approximate same scale; they are from approximately <25—x35, and are here reduced x 1/3. VOLUME 92, NUMBER 4 Fig. 3. Views of the tops, left, and bottoms, right, of the footplates (region of insertion into the oval window) of, from upper to lower, a trogon, kingfisher, and suboscine (same as in Fig. 2). SEM’s are from x45—x55, and are here reduced x%. 694 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 4. Zeiss differential interference contrast micrograph of several bundles of spermato- zoa from the Eastern Wood Pewee (Contopus virens). Approximately <700. microtubules are not arranged in a helical array with a mitochondrial com- ponent bound around the axoneme, and a mitochondrial component is not distinguishable. The undulating membrane characteristic in light microscopy of both oscine and suboscine sperm is due in the former case to a helically wound tripartite membrane, in the former to an undulating band of singlet microtubules that completely surround the axoneme. These data would not support a polyphyletic order Passeriformes, but would suggest that a com- mon ancestor had already evolved the sperm bundling and that the details of the ultrastructure underwent different changes in the two lines of evo- lution. It might be added that while there is currently no completely satisfactory definition of the Passeriformes, most have characteristic ‘‘aegithognathous’”’ palates and are generally similar osteologically. From the foregoing it now seems to me more probable that the order Passeriformes is monophyletic and the similar suboscine and alcediniform stapes evolved independently. This revised view would envisage a common ancestor for the suboscines and oscines before the late Cretaceous split of Gondwanaland, suboscines VOLUME 92, NUMBER 4 695 = ” ae b cok : fs SS we a ee Fig. 5. Electron micrograph of a transverse section through a bundle of spermatozoa of the Eastern Wood Pewee (Contopus virens) at a level posterior to the region of the nuclei. Ab- breviations are as follows: simt, singlet microtubules; pm, plasma membrane; df, dense fibers; and a, axoneme. Approximately <27,000. as a Southern Continent group, and oscines evolving in the Old World and not getting into South America before the rifting of the southern continents. Perhaps the most instructive insight into passerine relationships that the stapes has shown is that: (1) New World suboscines and Old World forms (Eurylaimidae, Pittidae and the Madagascan Philepittidae) are part of a monophyletic group, (2) oscines and suboscines are very distinctive groups, easily separable from one another, and (3) the two groups are separated by a broad and ancient evolutionary gulf. It may also be among the coraciiforms rather than piciforms that we need to search for the true sister group of the passerines. Acknowledgments I am indebted to Walter R. Brown, Mary-Jacque Mann, and Susann G. Braden who skillfully operated the scanning electron microscopes in the facility at the National Museum of Natural History, to Edward D. Salmon for taking the photos of the sperm bundles using differential interference contrast, and to Wilma Hanton for her help with the transmission EM. Sigrid James prepared the plates. I especially thank Storrs L. Olson, who encour- aged this undertaking; and R. L. Zusi, who criticized the manuscript and suggested the inclusion of the new sperm data. This study was undertaken 696 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON while the author was on a Kenan Leave from the University of North Car- olina. Literature Cited Feduccia, A. 1975. Morphology of the bony stapes (columella) in the Passeriformes and related groups: evolutionary implications.—Univ. Kansas Mus. Nat. Hist., Misc. Publ. No. 63. . 1977. A model for the evolution of perching birds.—Syst. Zool. 26:19-31. Henley, C., A. Feduccia, and D. P. Costello. 1978. Oscine spermatozoa: a light- and electron- microscopy study.—Condor 80:41-48. Wetmore, A. 1960. A classification for the birds of the World.—Smithsonian Misc. Coll. 139 (11): 1-37. Department of Zoology, University of North Carolina, Chapel Hill, North Carolina 27514 (and Research Associate, Department of Vertebrate Zoolo- gy, National Museum of Natural History, Smithsonian Institution, Wash- ington, D.C. 20560). PROC. BIOL. SOC. WASH. 92(4), 1979, pp. 697-718 A NEW GENUS OF WATER BEETLE FROM AUSTRAL SOUTH AMERICA (COLEOPTERA: HY DROPHILIDAE) Paul J. Spangler Abstract.—Descriptions and illustrations are given for the adult, larva, and pupa of a hydrobiine water beetle, Anticura flinti, new genus, new species, collected from Chile and Argentina. These forms are interpolated into existing keys. Pupation, habitat, collecting methods, and behavior are discussed. The adults are the first hydrophilids known to have dimorphic metathoracic wings and this condition is also discussed. A single female of this new genus was collected in 1974 from San Martin de los Andes, Argentina, by my colleague Dr. Oliver S. Flint, Jr. Although the temptation to describe the genus on the basis of the single female was strong, it seemed prudent to wait for more material; so the female was put aside until, hopefully, more specimens of both sexes could be obtained. In this instance patience was rewarded. The habitat of the single specimen collected by Flint was not known with certainty. Dr. Flint was collecting larvae and adults of Trichoptera from a small fast flowing stream when he collected the beetle, and it seemed prob- able that the specimen was found in piles of partly submerged driftwood or logjam habitats in the stream or at the margin of the stream. This type of habitat is similar to those in which we sometimes find Ametor latus (Horn) and A. scabratus (Horn) in the northwestern United States, Sperchopsis tessellatus Ziegler and Hydrobius melaenus (Germar) in the eastern United States, and Hydramara argentina Knisch in northwestern Argentina. There- fore, when the recent opportunity became available to go to Chile to par- ticipate in fieldwork in the mountains on the Chilean-Argentinian border, a search for this beetle was a first priority. Our work was centered in the region around Anticura in Puyehue National Park in Osorno Province. There we encountered dry weather which reduced stream flow in local streams and made collecting in them easy. The cold, cloudy, and misty mornings usually turned to warm, clear days by noon, and the surrounding mountains, volcanos, forests of Nothofagus, and the cold clean trout streams added to the esthetic pleasures of collecting in this area. A small stream, the Rio Anticura, flowed through the park and appeared to be a good one in which to search for the undescribed genus if it indeed occurred in this area. There- fore, I was delighted to discover a single hydrophilid larva in a logjam during 698 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON the first afternoon of collecting in the Rio Anticura. The larva had to be the undescribed larva of Hydramara argentina or, as it proved to be, the larva of the undescribed genus. The find was exciting, and it held promise of additional interesting specimens. On the second day of collecting we ex- plored the nearby Gol Gol River, and there Dr. Flint found a single adult of the undescribed genus. This adult was found, seemingly aestivating, un- der a rock in damp sand about 2 meters from a pool in the streambed. Knowing now that the undescribed genus occurred in the area, I returned to the Rio Anticura and began examining logjams in earnest. Finally my efforts were successful, and by the end of the day I had collected 34 adults and 15 larvae of the new genus. The size of a number of the larvae sug- gested that with intensive searching, a pupa might also be found. Additional collecting, mostly in the Rio Anticura, during the next 12 days provided a total of 73 adults and 26 larvae of the new genus but no pupae. Therefore, finding a single pupa with its shed larval skin during my last day of collecting was especially rewarding. During the entire collecting period a search was made for eggs or egg cases, but none was found; probably it was too late in the breeding season. All 73 specimens of the new genus found in the Rio Anticura were collected in the river behind the park headquarters at Anti- cura. Hydrobiinae Anticura, new genus Body form (Fig. 1) broadly ovate, moderately convex. Head (Figs. 2, 3, 4) with clypeus expanded shelflike in front of eyes and strongly emarginate medially on anterior margin. Labrum similarly emarginate medially and an- terior margin barely visible beneath clypeus. Eyes viewed from above ovoid. Antenna (Fig. 4) 9 segmented; 2 basal, 3 intermediate, 1 cupule, and 3 club segments; only club segments pubescent. Maxillary palpus (Fig. 4) moderately long, robust, and 4 segmented; basal segment very short; second (pseudobasal) segment longest, subequal to ultimate and penultimate com- bined; ultimate segment subequal to penultimate segment. Prosternum not carinate medially, terminating posteromedially in a bisinuate border, and lacking a prosternal process. Mesosternum (Fig. 5) with short, feeble lon- gitudinal carina between apical halves of mesocoxae and behind a low, transverse arcuate ridge across posterior third of mesosternum. Metaster- num apically with short narrow carina between and slightly behind posterior halves of mesocoxae (Fig. 5); disc of metasternum higher than sides and bearing a narrow, longitudinal, glabrous area on midline on posterior fourth; behind glabrous area, metasternum terminates in an elongate triangular sclerite which extends part way between metacoxae on midline. Elytron with sutural stria; broad, moderately convex; shining; bearing 10 distinct VOLUME 92, NUMBER 4 699 rows of very coarse punctures, a partial eleventh row laterally; epipleura wide and horizontal basally, narrowing abruptly and becoming almost ver- tical opposite third abdominal sternum. Scutellum an elongate triangle. Front and middle femora, except about apical eighth, densely covered on all surfaces with hydrofuge pubescence. Hind femur lacks hydrofuge pu- bescence on ventral surface but bears fine sparse punctures, each puncture bearing a short seta. Hind tibia not arcuate, without fringe of long natatory hairs. Tarsal formula 5-5-5. Middle and hind tarsi with first segment slightly shorter than second segment (Fig. 6). Type-species of the genus.—Anticura flinti, new species. Etymology.—Anticura, from the name of the lovely river from which adults, larvae, and pupa were collected. Gender: feminine. This new genus keys to the subfamily Hydrobiinae in our latest keys to the higher categories of the Hydrophilidae, e.g., Crowson (1955) and Leech (1956). However, to compare this new taxon with previously described gen- era it is necessary to use d’Orchymont’s (1942) revision of the taxa consid- ered to belong to the tribe Hydrobiini at that time. In his study d’Orchymont provided a key to two divisions to which he referred as subtribes, i.e., the Hydrobiae and the Helocharae. The new genus, Anticura, belongs to d’Orchymont’s subtribe Hydrobiae which corresponds roughly to our pres- ent concept of the Hydrobiinae. However, in Anticura the maxillary palpi are slightly longer than the antennae contrary to d’Orchymont’s statement that the maxillary palpi are shorter than or as long as the antennae. Although some other characters used by d’Orchymont also disagree with those pres- ent in this new genus, those characters were included by him to accom- modate the genera Laccobius Erichson, Beralitra d’Orchymont, and Oocy- clus Sharp which do not belong in the Hydrobinae. After comparing adults and larvae of Anticura with adults and larvae of Hydrobius Leach, Hydra- mara Knisch, Sperchopsis LeConte, and Ametor Semenov (as well as with the rygmodine genera Cylorygmus d’Orchymont and Rygmodus White) and until a new key to world genera of the Hydrophilidae is available, I have interpolated this new genus into d’Orchymont’s key to his Hydrobiae. The new genus Anticura keys to couplet 7 in d’Orchymont’s (1942:6) key to the genera of the subtribe Hydrobiae. The following modified couplets will separate Anticura from the other genera keying to d’Orchymont’s cou- plet 7. 7a. Posterior femur without dense hydrofuge pubescence on ventral surface; middle femur with or without hydrofuge pubescence .... 7b — Posterior femur with dense hydrofuge pubescence on ventral sur- face; middle femur similarly pubescent ....................2000- 7c 7b. Middle femur pubescent; deeply emarginate labrum hidden under deeply emarginate clypeus; Chile and Argentina . Anticura, new genus 700 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON — Middle femur not pubescent; labrum completely visible in front of elypeus; Australia joe citeorsobmeuiier uysteree ce vel petri ae Hybogralius 7c. Anterior edge of pronotum shallowly emarginate; form more de- pressed; northwestern United States; China to Nepal ........... 7d — Anterior edge of pronotum deeply emarginate; anterolateral angles of pronotum extending almost to anterior margins of eyes; form very convex; eastern United States and Southeastern Canada.... aie bo seaieenen yey dia ti een taba A aeds iaylts bea as Dea seer Sperchopsis 7d. Elytral intervals smooth between the dual punctures. Epipleuron not truly horizontal at base, rapidly bent and becoming more ver- tical; the metasternum and abdomen hidden in elytral cavity; me- sosternal process moderately developed and projecting; China ... Ee ne er er see Se BNE Sb 5 ici boc Hydrocassis — Elytral intervals never smooth, densely and uniformly scabrous or granulose; epipleuron nearly horizontal at base and becoming more so apically; metasternum and abdomen only slightly hidden in ely- tral cavity; mesosternum without projecting process or at most only slightly swollen between mid coxae; northwestern United States; China, to-Nepal «.. ..0.ce cient se BR eee et ee Ametor Anticura flinti, new species Figs. 1—25 Holotype male.—Body form (Fig. 1) broadly ovate. Length 6.5 mm; great- est width 4.0 mm, slightly behind midlength. Color above piceous except thinner margins of elytra dark reddish brown. Ventral surface piceous ex- cept as follows: antennae and apices of palpi light reddish brown; legs dark reddish brown. Entire venter (except legs) behind mentum densely covered with dark reddish brown hydrofuge pubescence. Head shining but densely covered with coarse and fine intermixed punc- tures; coarse punctures about five times as large as fine punctures, becoming coarser and sometimes confluent especially between and behind eyes and to base of head; base sharply recessed for reception of anterior edge of pronotum; recessed area covered with hydrofuge pubescence continuous with that behind eyes and on ventral surface of head. Clypeus greatly ex- panded and shelflike in front of eyes, covering all but narrow anterior margin of labrum; strongly broadly emarginate apicomedially (Figs. 2, 3, 4). Labrum mostly hidden but narrow exposed portion strongly emarginate conforming to emargination of clypeus; bearing many dense punctures and short, stout setae on anterior margin. Ventral surface of head with microsculpture and pubescence except mentum, maxillae, and other appendages smooth; stipes of maxillae glabrous and finely sparsely punctate; mentum (Fig. 7) broadly and moderately deeply emarginate apicomedially and smooth except a few 701 VOLUME 92, NUMBER 4 Anticura flinti, n. gen., n. sp., d holotype, habitus view. Fig. 1. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Figs. 2-3. Anticura flinti, n. sp., head: 2, Anterior view, 54x; 3, Lateral view, 57x. VOLUME 92, NUMBER 4 703 fine lateral punctures. Antenna (Fig. 4) 9 segmented; 2 basals, 3 interme- diate, 1 cupule, and 3 club segments. Maxillary palpus (Fig. 4) 4 segmented; basal segment very short; second segment longest, about equal to third and fourth segments combined; penultimate segment slightly longer than ulti- mate segment, wider apically. Labial palpus small; 3 segmented; first seg- ment short, about a fourth as long as second segment; second segment about as long as third segment but broader and bearing a tuft of long yel- lowish-brown setae along anterior edge; third segment almost parallel sided, only slightly wider apically. Pronotum slightly more than twice as wide as long; strongly arcuate lat- erally; punctures much coarser than those on head and without fine inter- mixed punctures; punctures more dense laterally than on discal area; an- terior, lateral, and posterolateral third of posterior edges finely margined; anterolateral and posterolateral angles strongly rounded; lateral edge of hy- popleura with coarse, closely spaced punctures. Scutellum an elongate triangle with base about three-fourths as long as sides; bearing 2 punctures. Elytron with sutural stria; broad, moderately convex; shining; bearing 10 distinct striate rows of very coarse punctures, a partial eleventh row later- ally; intervals impunctate; epipleura wide and horizontal basally, narrowing abruptly and becoming almost vertical opposite third abdominal sternum. Prosternum almost flat medially, terminating posteromedially in an ar- cuate border and lacking a prosternal process. Mesosternum (Fig. 5) with short, feeble, longitudinal carina between apical halves of mesocoxae and behind a low, transverse, arcuate ridge across posterior third of mesoster- num. Metasternum (Fig. 5) apically with short, narrow carina between and slightly behind posterior halves of mesocoxae; discal area higher than sides and bearing a narrow, longitudinal glabrous area on midline on posterior fourth. Behind glabrous area, metasternum terminates in an elongate tri- angular sclerite which extends part way between metacoxae on midline. Front and middle legs with femur densely covered with hydrofuge pu- bescence on all surfaces except apical eighth glabrous. Hind femora lack hydrofuge pubescence on ventral surface but bear fine sparse punctures, each puncture bearing a short seta. Hind tibia not arcuate, without fringe of long natatory hairs. Tarsal formula 5-5-5. Middle and hind tarsi with first segment only slightly shorter than the second segment (Fig. 6). Foreleg with _segments 1 to 4 about equal in length; last segment robust, swollen apically, and subequal to segments 1 to 4 combined. Midleg with segments 1, 3, and 4 equal in length; first segment about two-thirds as long as second segment; last segment robust, swollen apically, and about four-fifths as long as seg- ments 1 to 4 combined; all tarsal claws long and robust. Abdominal sterna covered with hydrofuge pubescence. Last visible ab- dominal sternum rounded, not emarginate apicomedially. Genitalia as illustrated (Figs. 8, 9). PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Figs. 4-5. Anticura flinti, n. sp.: 4, Antenna and maxillary palpus, 75x; 5, Mesosternum, mesocoxae, and apex of metasternum, 110. x LY @) nN Oo < ie =) n — i 0) > nel io} (0) an ~ x Ww ™~ a =) N fw 9] _ uo) = a0) N-} a, N q = i S 8 x = AS) = = x 3 7) oD — ply VOLUME 92, NUMBER 4 706 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Figs. 8-9. Anticura flinti, n. sp., holotype, male genitalia: 8, Ventral view; 9, Dorsal view. Allotype.—Similar to male; no external sexual differences could be found. Variations.—The only variations evident among the specimens in the type-material are the usual color and size differences. Teneral specimens are light brown to dark reddish brown instead of piceous as in older spec- imens. Also, the specimens vary in size as follows. Male: length, 5.0 to 6.1 mm; width, 3.4 to 4.0 mm. Female: length, 6.0 to 6.6 mm; width, 3.8 to 4.5 mm. Etymology.—I am pleased to dedicate this new species to my friend Dr. Oliver S. Flint, Jr., who collected the first specimen of this species and who has collected many other aquatic Coleoptera for me during his many field excursions. Type-data.—Holotype male: CHILE: Osorno Province, Anticura, Puye- hue National Park, in Rio Anticura, 2 Feb. 1978, Paul J. Spangler, USNM Type No. 75765, deposited in the National Museum of Natural History, Smithsonian Institution, Washington, D.C. Allotype: Same data as holo- type. VOLUME 92, NUMBER 4 707 Paratypes: ARGENTINA: Neuquen Province, San Martin de los Andes, Arroyo Rosales, 22 Jan. 1974, O. S. Flint, Jr., 1 2. CHILE: Same data as holotype, 20 d¢, 12 22; Osorno Province, Anticura, Puyehue National Park, in Rio Gol Gol, 1 Feb. 1978, Oliver S. Flint, Jr., 1 2; Osorno Province, Puyehue National Park, Anticura, in Rio Anticura, 3 Feb. 1978, Paul J. Spangler, 6 dd, 11 2 2; Osorno Province, Puyehue National Park, Aguas Calientes, in Rio Chanlefu, 8 Feb. 1978, Paul J. Spangler, 2 2 2; Osorno Province, Puyehue National Park, Anticura, in Rio Anticura, 11 Feb. 1978, Paul J. Spangler, 14 65,622. Paratypes will be deposited in the following entomological collections: American Museum of Natural History, New York; British Museum (Natural History), London; California Academy of Sciences, San Francisco; Cana- dian National Collection, Ottawa; Institut flr Pflanzenschutzforschung Zweigstelle, Eberswalde; Museo Argentino de Ciencias Naturales, **Ber- nardino Rivadavia,’’ Buenos Aires; Museo National de Historia Natural, Santiago; Museum fur Naturkunde, East Berlin; Museum National d’Histoire Naturelle, Paris; Institut royal des Sciences naturelles de Bel- gigue, Brussels; and Zoologische Sammlung Bayerischen Staates, Mun- chen. Description of Third-instar larva Figures 10-19 Body elongate (Fig. 10). Total length, 14.8 mm; width of prothorax 1.7 mm. Color of integument grayish brown. Sclerotized head capsule, thoracic sclerites, spiracles, and legs yellowish brown. Integument covered with as- perities whose apices are oriented posteriorly but otherwise disarrayed. Head (Fig. 12) quadrangular, 1.6 mm wide, 1.2 mm from labroclypeus to occipital foramen. Frontoclypeal suture feebly indicated. Frontal sutures united near base of head forming an epicranial suture. Frons sagittate. Cer- vical sclerites present, rectangular. Ventral surface of head with numerous setae laterally, glabrous medially except a short row of 3 slender setae basolaterally on each side of midline; with 2 deep, approximate, posterior tentorial pits behind gula. Labroclypeus (Fig. 14) prominent, almost symmetrical; with 2 large me- dial teeth, left tooth slightly shorter than right tooth; 4 stout setae present, 1 on each side of each tooth. Anterolateral projections of epistoma shorter than teeth, left projection slightly shorter than right; both projections sub- angular anteromedially, broadly rounded anterolaterally, and both bearing a few short setae on anterior margins. Ocular areas each with groups of 6 distinct ocelli arranged in an ellipse; anterior 3 ocelli larger and close to each other; posterior 3 ocelli smaller, ventrolateral one smallest and separated from posteromedial two. 708 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 10. Anticura flinti, n. sp., larva, habitus view. VOLUME 92, NUMBER 4 709 Antenna (Fig. 19) 3 segmented, short, cylindrical, shorter than stipes; first segment longest, about one-third longer than penultimate segment; penul- timate segment with small apicolateral tubercle and a long slender apico- medial seta; ultimate segment small and slender, about one-fifth length of penultimate segment and bearing a distinct, slender, distal seta and a minute apicolateral appendage. Mandible (Figs. 11, 13) symmetrical, prominent, stout, sharply pointed apically; each with 2 large well-defined inner teeth and 1 large distal tooth; molar area rounded. Maxilla (Fig. 16) with stipes stout, elongate, tapering distally, bearing a row of 12 or 13 stout setae on inner margin. Palpiger segmentlike, with slender sclerotized appendage on apicomedial angle about one-third as long as palpiger; bearing 1 long seta medially below appendage; appendage of palpiger bearing an apical seta. Palpus 3 segmented, tapering distally; first segment short, about half as long and about half as wide as palpiger; pen- ultimate segment longest, twice as long as basal segment and bearing | lateral seta at about apical third; ultimate segment conical, about half as long as penultimate segment and bearing 1 basomedial seta. Labium (Fig. 17) extending about to midlength of stipes. Penultimate seg- ment of palpus short and broad; ultimate segment not quite twice as long as penultimate segment. Ligula distinct, about a third longer than penulti- mate palpal segment and bearing a basal seta at each lateral corner. Mentum subquadrangular, glabrous except 1 lateral and 1 ventrolateral seta, wider than submentum, arcuate laterally, narrowing posteriorly; dorsal surface spinose; with numerous setae on anterior edge; ventral surface glabrous except for apicolateral setae. Gula pentagonal, rounded posteromedially. Prothorax with sides nearly straight but diverging posteriorly; postero- lateral angles broadly rounded, with a few short setae; anterolateral angles each with a few long setae and a few short setae; sagittal line present. Prosternal sclerite large, subrectangular, with no indication of sagittal line. Mesothorax wider than prothorax and about half as long (measured on midline) as prothorax; with 2 small, very narrow, straplike, anterior sclerites and 2 large, subrectangular mesotergal sclerites; lateral margins each with a prominent spiracular tubercle which is followed by a sparsely setiferous lobe; sagittal line present. Metathorax slightly wider than and about as long as mesothorax; anterior metatergal sclerites transverse, larger than posterior sclerites, irregularly rectangular; posterior sclerites small, narrow; sagittal line present. Legs four segmented; procoxae large, separated by about length of a trochanter; trochanter as long as femur (viewed ventrally); femur about as long as tibiotarsus; tarsal claw single, ventrally bearing 2 large robust setae at basal third. 710 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Ay M4 | AA 4 Ui A ; VOLUME 92, NUMBER 4 711 Abdomen of 8 distinct segments, ninth and tenth segments reduced; terga similar to each other and separated by an intersegmental membrane. True segmentation obscured by additional transverse folds on segments, segmen- tal folds continued onto sternum. Each segment with 4 folds; anterior fold with 4 small setose tubercles; second and third folds without tubercles; fourth fold with 2 large medial tubercles and 2 smaller lateral tubercles. All tubercles are densely covered with asperities. A large spiracular tubercle present on each side of segments | through 7. Epipleurites and hypopleurites prominently lobed. Eighth tergum represented by superior valve of stigmatic atrium which bears 2 large sclerites (Fig. 15), beneath which lies the eighth pair of abdominal spiracles. Ninth tergum trilobed. Middle lobe large; with 2 short, stout setae, 1 on each side of median line on caudal margin. Lateral lobes small, about a fourth as wide as median lobe; each bearing 2 elongate, slender setae on posterior margin. Specimens examined (30): All from CHILE, Osorno Province, Puyehue National Park: Anticura, in Rio Anticura: 1 Feb. 1978, 2 larvae; 2 Feb. 1978, 15 larvae; 3 Feb. 1978, 4 larvae; 11 Feb. 1978, 6 larvae. Aguas Calientes, in Rio Chanlefu: 7 Feb. 1978, 1 larva; 8 Feb. 1978, 2 larvae. The larva of Anticura flinti traces to couplet 23 in Bertrand’s (1972:263) generic key to hydrophilid larvae. However, Anticura may be distinguished from the two choices available at this point by inserting the following mod- ified couplet there. Da waAbrOclypeus) with 2 teethhy .26 69. cee wwe ces Anticura, new genus ay I ADLOCIYPECUIS WAUME4IOR SD LOCUM osu a, so ree ke « he eens seas 23b y8be Labroclypeus with 4 teeth...) . 6. ss sects ete vee ee eae 24 ee ADKOCIVPEUS WAlMy Sr UCC MMe. fs cick. oisiep raid 6, ou0%G save chake oS S7s. 5 dye oO 26 Description of Pupa Figs. 20-21 Total length 10.4 mm, greatest width 4.5 mm. Color white except styli and cerci light yellowish brown. Glabrous except for setae and styli de- scribed below. Head with 2 supraorbital styli above each eye and 1 (possibly 2) short seta on each side of midline on vertex. Pronotum with 22 styli as follows: 3 on each anterolateral angle, 2 on each side of median line on anterior margin, 3 on each posterolateral angle, 2 on each side of median line at posterior margin and 1 on each side of median line on disc. Mesonotum with 2 styli, 1 on each side of scutellum. Meta- notum with 2 styli, 1 on each side of midline. <— Figs. 11-19. Anticura flinti, n. sp., larva: 11, Left mandible; 12, Head; 13, Right mandible; 14, Labroclypeus; 15, Stigmatic atrium; 16, Maxilla; 17, Labium, dorsal view; 18, Labium, ventral view; 19, Antenna. 712 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON la 20 21 Figs. 20-21. Anticura flinti, n. sp., pupa: 20, Dorsal view; 21, Ventral view. Abdomen with 4 styli on first segment; second to seventh segments each with | pair of pleural and 6 tergal styli (total 8) arranged as follows: 1 pleural stylus lateral to each spiracle, 1 stylus behind each abdominal spiracle, and 2 styli between each spiracle and midline. Segment 8 with 2 styli on posterior margin, 1 on each side of midline. Segment 9 with 2 cerci slightly longer than basal width of segment 9, each cercus terminating in a single stout seta. First to seventh abdominal segments each with a pair of spiracles; those on segments | and 7 reduced, especially so on the seventh segment. Antennae and femora extending outward at right angles from body axis. Tibiae of all legs folded back against their respective femora. Tarsi turned backward and parallel with body axis, those of front and middle legs widely separated, hind tarsi approximate. The partially developed parameres and median lobe of the male genitalia visible at the apex of the abdomen show that the pupa described above is a male. VOLUME 92, NUMBER 4 713 Specimen examined: CHILE, Osorno Province, Puyehue National Park, Anticura, in Rio Anticura, 11 Feb. 1978, 1 pupa. The pupa (Figs. 20, 21) of Anticura keys to couplet 9 (i.e., Ametor) in Bertrand’s (1972) key to hydrophilid pupae, but it may be distinguished from Ametor immediately because the pupa of Anticura has 22 instead of 20 pronotal styli and 1 pair of pleural and 6 tergal instead of 1 pair of pleural and 4 tergal styli as are present on Ametor. Pupation Pupation occurs in loam in pupal chambers out of water as is character- istic for most hydrophilids whose biologies are known. The single pupa I found was in a chamber in loam mixed with a few leaves caught between two rocks, i.e., not in a typical logjam habitat. Several teneral and very lightly pigmented adults were found in loam in one logjam. Habitat Almost all of the specimens of Anticura flinti were collected in logjams (Figs. 22, 23) where:logs, sticks, twigs, leaves, as well as loam were de- posited and compacted by the current. Occasionally grasses or other plants were growing in this compacted debris, their roots binding the debris in a matlike fashion. All of the larvae were found on the sticks or logs in the logjams. Adult beetles were found on the logs or in cracks and crevices in the logs, among the compacted leaves and roots, and occasionally on rocks present under the matted roots. Several adults were found along the under- side of a single large log, one end of which was shallowly imbedded in the loamy stream bank. Because many of these logjams were situated where there was considerable current, some beetles were washed away as their habitat was necessarily destroyed in the process of searching for them. One adult collected by Dr. Flint was found several feet from water under a rock on a sandy shore in a branch of the Gol Gol River. That beetle apparently retreated to the damp sand there as the water level in the stream dropped drastically leaving only small isolated pools in the streambed. During the day when I collected the specimens of A. flinti, the air tem- perature varied from 54°F to 71°F. The water temperature varied from 44°F to 48°F. The rivers in this area flow through rocks of volcanic origin. Col- orimetric water testing methods for pH and hardness indicated the follow- ing: pH 5S and 0 grains per gallon for the Rio Anticura. Specimens were collected at altitudes ranging from 330 to 700 meters. Behavior Larvae found on sticks moved slowly but kept their jaws open and head 714 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON a” , PA. / J nn Se ss s Se ee ae j f = ~~ 4 , Z 4 i St AE ae %/, ae « h ¥ ¢ ‘x , a ~3 ao abt. a ato “ : R 1 gee, mn OPS Bo. Pn oa es = be : a Figs. 22-23. Anticura flinti, n. sp., habitat: 22, Rio Anticura at type-locality; 23, Logjam biotope. moving back and forth from side to side as they actively searched for prey. Larvae seemed to wash loose rather easily from their footholds in contrast to the adults which were often in crevices in logs or in the matted debris and not easily dislodged. However, those adult specimens disrupted from smaller sticks or under the edges of mats often dropped off into the water where the current caught them and whisked them away. When disrupted in the logjam adults usually feigned death and remained motionless for varying lengths of time. When they became active again they moved rather slowly and in short, sometimes jerky movements. Adults that were dislodged into the stream or that were placed in small artificial pools of water floated about inactively until they bumped against an object or the shoreline, then they grasped any convenient foothold and tried to crawl away and hide or burrow into coarse sand. None attempted to swim. VOLUME 92, NUMBER 4 715 Collecting Methods At first I pulled logs, sticks, rocks, matted roots, etc., slowly out of the logjam while watching for larvae and adults. However, after losing several specimens of this rare beetle into the current I moved only the large logs and then gathered all of the debris or as much as possible in one or a few handfuls and placed this material on a square meter of nylon cloth on the shore. I then collected the exposed larvae and adults immediately. As the sun heated and dried the debris, other specimens became active and were then easily seen moving and were collected. Wing Dimorphism Although the majority of insects have fully developed wings and are ca- pable of flight, some are brachypterous and unable to fly. The ability or inability to fly obviously affects the vagility and survival of the species depending upon its habitat. In aquatic beetles, most taxa have normal and functional wings. Therefore, it is very interesting to find exceptions to the usual condition. Little information is available on wing dimorphism in the family Hydrophilidae because, as Jackson (1928) reported in a brief sum- mary, wing dimorphism seems to be very rare in this family. In an interesting study of wing atrophy in the family Carabidae, Darlington (1936) reported that he had examined a considerable number of aquatic and semiaquatic beetles in the families Amphizoidae, Omophronidae, Haliplidae, Dytiscidae, Gyrinidae, Hydrophilidae, Psephenidae, and Heteroceridae without finding a single eastern American species without normal flight wings except the dytiscid Ilybiosoma bifarius (Kirby). Reporting on flight in water beetles, Jackson (1952) noted that in the Hydrophilidae (sensu lato), the species Helophorus ytenensis Sharp, Hydraena minutissima Stephens, and Hy- draena pygmae Waterhouse have abbreviated wings. Since Jackson’s paper appeared in 1952, hydraenid beetles have been removed from the family Hydrophilidae and have been placed in their separate family, the Hydraen- idae. Furthermore, Helophorus is variously placed in a separate family or subfamily by some authors. Therefore, the only uncontested species with reduced flight wings presently included in the Hydrophilidae (sensu stricto) seems to be Coeloctenus seriatus Balfour-Browne from Africa. When this species from the shores of Lake Tanganyika was described by Balfour- Browne (1939) he reported that in his specimens ‘‘the flight wings are re- duced to mere functionless slips.’’ Balfour-Browne further stated that ‘‘the reduction of the wings is a feature of great interest, conceivably correlated with the habitat, since reduced wings are less likely to impede the insect in the event of involuntary raising of the elytra in rough water.’’ Therefore, the discovery of both normal (Fig. 24) and reduced (Fig. 25) wings in An- ticura flinti, represents the second species in the Hydrophilidae (sensu stric- 716 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON to) with flightless specimens and apparently the first with both normal and reduced wings in the same species. The presence of normal and reduced flight wings in Anticura flinti, was discovered by chance because having seen the flight wings extended beyond the elytra in some specimens I had preserved in alcohol, I assumed they all had normal flight wings. However, after having pointed and labeled many of my specimens I decided to remove for study a flight wing from a specimen with a broken elytron. Upon lifting both elytra I discovered only the vestigial wing on both sides of the metanotum. Consequently, I examined the 36 unmounted specimens and found that 31 had reduced and flightless wings and only 5 had normally developed wings. Of the 31 with reduced wings, 20 were males and 11 were females. Of the 5 specimens with normal wings, 3 were males and 2 were females. Three of these 5 winged specimens were teneral and very lightly pigmented suggesting that they had enclosed prob- ably within the previous 24 to 48 hours. Unfortunately, I had only 5 beetles remaining in alcohol with normal wings and preferring to keep them as paratypes I did not dissect any to try to decide whether their flight muscles were developed sufficiently for flight. For beetles living in mountain streams like the Rio Anticura, Rio Chan- lefu, and Rio Gol Gol, the ability to fly would be a definite advantage be- cause the polished rocks and numerous piles of huge Nothofagus and other tree trunks in these streams (Fig. 22) attest to the apparently common scour- ing of these streambeds by high water. The distribution of the hydrofuge pubescence present on A. flinti and personal observations verify that this Species carries a bubble of air over most of its ventral surface and in the air pocket beneath its elytra when it submerges as is usual for other hydro- philids. Therefore, unlike elmids which can obtain their air by means of plastron respiration and need not surface, A. flinti must surface periodically to replenish its air supply. This need to surface for air amidst the physical force of the fast current during floods would be detrimental to the survival of the flightless specimens of this species. Considering the above circumstances it seems probable that normal- winged individuals, although low in numbers, are capable of flying and are the main source of dispersal of this species from stream to stream. After a population becomes established in a stream the species probably spreads downstream by drifting of the larvae and flightless specimens and, certainly to some extent, by flight by the winged specimens. Further studies of wing dimorphism, generations per year, dispersal, genetics, and distribution in this species should be very interesting and illuminating. Acknowledgments I am grateful to Luis E. Pena G. who was responsible for organizing the VOLUME 92, NUMBER 4 717 2) Figs. 24-25. Anticura flinti, n. sp., metathoracic wing: 24, Normal wing; 25, Vestigial wing. cooperative fieldwork between the J. I. Molina Institute of Studies and publications of Chile and the Smithsonian Institution during which this new genus was collected. Furthermore, his knowledge of Chile and its flora and fauna, his experience in fieldwork, his culinary delights, and his keen sense of humor made him an ideal leader for the project. I am also pleased to acknowledge the financial support received from the J. I. Molina Institute and the gracious hospitality and friendship extended to me by members of the Institute who were able to meet with us and participate in some of the fieldwork. In addition, I thank the Smithsonian Institution administrators for a Fluid Research Grant which contributed to my participation in this rewarding fieldwork in Chile. My thanks are extended also to the following individuals for their assis- tance with this study. Dr. Oliver S. Flint, Jr., who collected the first spec- imen of this new genus; Ms Mary Jacque Mann, Smithsonian Institution scanning electron microscopist, for the micrographs; and Mr. Michael Druckenbrod, Smithsonian Institution staff artist, for preparing the illustra- tions. Literature Cited Balfour-Browne, J. 1939. Contribution to the study of Palpicornia. II.—Ent. Mo. Mag. 75:1- 8. Bertrand, H. P. I. 1972. Larves et Nymphes des Coléoptéres Aquatiques du Globe. 804 pp., 561 figs.—Abbeville, France: F. Paillart. Crowson, R. A. 1955. The Natural Classification of the Families of Coleoptera. 187 pp., 213 figs.—London: Nathaniel Lloyd and Company, Ltd. 718 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Darlington, P. J., Jr. 1936. Variation and Atrophy of Flying Wings of Some Carabid Beetles.— Ann. Ent. Soc. America 29(1):136—-179, 3 pls. , Jackson, D. J. 1928. The Inheritance of Long and Short Wings in the Weevil (Sitona hispidula) with a Discussion of Wing Reduction among Beetles.—Trans. Roy. Soc. Edinburgh 55:665—735. . 1952. Observations on the Capacity for Flight of Water Beetles.—Proc. Roy. Ent. Soc. London (A) 27:57-70. Leech, H. B. 1956. Aquatic Coleoptera, Chapter 13, in: Aquatic Insects of California with keys to North American genera and California species. (edited by R. L. Usinger). Pages 293-371.—Berkeley, California: University of California Press. d’Orchymont, A. 1942. Contribution a L’étude de la tribu Hydrobiini Bedel, spécialement de sa sous-tribu Hydrobiae (Palpicornia-Hydrophilidae).—Mem. Mus. Roy. Hist. Nat. Bel- gique, ser. 2, fasc. 24:1-68, 4 figs. Smithsonian Institution, Washington, D.C. 20560. PROC. BIOL. SOC. WASH. 92(4), 1979, pp. 719-723 NEOTROPICAL MONOGENOIDEA, 1. OLIGAPTA KRUIDENIERI N. SP. (AXINIDAE: INDOCOTYLINAE) FROM THYRINOPS PACHYLEPIS (GUNTHER) IN EL SALVADOR John W. Crane, Delane C. Kritsky, and Robert J. Kayton Abstract.—Oligapta kruidenieri n. sp. (Axinidae: Indocotylinae) is de- scribed from the gills of Thyrinops pachylepis (Gunther) from an estuary located 13°15’N; 88°40’W (north shore of Peninsula de San Juan del Gozo, El Salvador). This species is distinguished from other species of the genus by having only 3-4 testes, an unarmed cirrus, vitellaria extending to phar- ynx, and by being less than 1 mm in length. The generic diagnosis of Oli- gapta Unnithan, 1957, is emended. Thus far, only 2 species of Oligapta have been described, both from the gills of marine fishes of the Eastern Hemisphere. Unnithan (1957) proposed the genus for O. oligapta collected from Hemiramphus georgeii (Val.) from Mandapom Camp, India. Young (1968) described O. manteri from gar pike, Hemiramphus sp., from Noumea, New Caledonia. As part of a survey of the monogenoid fauna of El Salvador (Central America) during 1976, 5 spec- imens of an undescribed species of Oligapta were recovered from the gills of 2 Thyrinops pachylepis (Gunther) collected from an estuary located 13°15'’N; 88°40’W (north shore of Peninsula de San Juan del Gozo). This species is described herein as O. kruidenieri (respectfully named for Dr. F. J. Kruidenier, University of Illinois, Urbana). Hosts were collected by dip net and immediately placed into a 1:4,000 mixture of formalin and seawater. After 1 hour, sufficient formalin was added to the containers to make a 5% solution. Monogenoids were removed directly from the gills and stored in 10% formalin. Techniques for the prep- aration and study of the helminths were those of Kritsky et al. (1972). Mea- surements, in wm, were made according to the procedures of Dillon and Hargis (1965). Figures were prepared with the aid of a camera lucida or microprojector. Type specimens were deposited in the U.S. National Mu- seum of Natural History Helminthological Collection (USNM 75211, holo- type; 75212, paratypes); and the University of Nebraska State Museum (No. 20975, paratype). Fish hosts were deposited in the American Museum of Natural History (No. 38203). 720 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Oligapta kruidenieri, new species Figs. 1-5 Description (based on 1 immature and 4 adult specimens; adults mea- sured).—Length 515 (484-586), greatest width 187 (142-224) near level of reproductive organs. Body lanceolate, stout. Tegument thick with numerous minute folds. Anterior cephalic margin with 3 (1 terminal) incipient lobes; well-developed glandular area anterior to prohaptoral suckers. Prohaptoral suckers 2, lying in posterolateral wall of buccal funnel; sucker subovate, 29 (26-33) by 21 (18-24), with conspicuous septum. Haptor 187 (158-209) long, 204 (155-245) wide, diagonal, armed with 8 sessile clamps and 2 pairs of anchors; clamp formation suppressed in adult. Clamps similar; sclerites del- icate, fragile; center piece with simple dorsal and T-shaped ventral termi- nations; dorsal loop sclerite incipient; clamp 67 (53-84) wide, 55 (47-62) long. Anchors dissimilar, situated dorsal to the 4th, 5th, and 6th clamps. Median anchor with lunate blade, elongate shaft originating from base of blade; anchor 37-38 long. Lateral anchor sickle shaped with elongate deep root; anchor 36-37 long. Pharynx ovate, 16 (12—21) wide, lying immediately posterior to buccal funnel; esophagus elongate, simple; crura obscured. Testes 3—4, irregular; each 19 (17-21) in diameter. Seminal vesicle, prostates absent; vas deferens obscured. Genital atrium 22 (19-24) in diameter, mid- ventral at level of bifurcation of gut, armed with 26-31 spines disposed into 2 tiers; spines 6—7 long, with recurved point, simple base. Ovary J-shaped, lying dextral to midsagittal plane. Seminal receptacle immediately anterior to ovary; oviduct, uterus delicate; odtype sinistral to posterior portion of Ovary; vitelline reservoir Y-shaped, with long sinistral branch. Vitellaria dense throughout trunk except absent in region of gonads. Vagina dextral, with distal sclerotized cone; cone 27 (23-30) long. Genitointestinal canal, excretory system, nervous system indistinct; eyes absent. Remarks.—Oligapta kruidenieri n. sp. is the only species in the genus possessing an unarmed cirrus and simple ventral clamp sclerites. Although some of the clamps in our specimens showed fractures of the ventral and dorsal sclerites, we consider these to be artifacts caused by coverslip pres- sure. The fractures were variable and apparently random in position in the 4 available adult specimens. The new species appears most closely related to O. oligapta as shown by the morphology and position of the sclerotized vagina, and the structure of the genital atrium. It differs from this species and O. manteri by having an unarmed cirrus, only 3 or 4 testes (25-30 in O. oligapta; up to 57 in O. manteri), vitellaria extending anterior to level of pharynx (vitellaria coex- tensive with crura in O. oligapta and O. manteri), and by being less than 1 mm in length. The new species is the first of the genus reported from the Western Hemisphere. 721 VOLUME 92, NUMBER 4 Oligapta kruidenieri: 1, Whole mount of holotype (ventral); 2, Sclerotized va- Figs. 1-5. ginal cone; 3, Genital atrium and cirrus; 4, Anchors; 5, Clamp (ventral). W22 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Oligapta Unnithan, 1957 Emended generic diagnosis.—Axinidae, Indocotylinae. Body divisible into cephalic region, trunk, haptor. Haptor with diagonal row of 8 sessile, Microcotyle-like clamps; 2 pairs of anchors. Prohaptoral suckers 2; pharynx small; esophagus without lateral diverticula; crura blind, terminating ante- rior to haptor, with or without diverticula. Testes few to numerous, post- ovarian, intercecal. Cirrus armed or unarmed; genital atrium midventral, armed, lying near level of bifurcation of gut. Ovary elongate, intercecal, with ends directed anteriorly. Vagina unarmed, dextral, posterior to genital atrium, with a distal sclerotized cone or peg. Seminal receptacle immedi- ately anterior to distal portion of ovary. Vitellaria extensive in trunk. Par- asites of marine fishes. Type-species.—O. oligapta Unnithan, 1957, from Hemiramphus georgeii (Val.), Mandapom Camp, India. Remarks.—The emendation of the generic diagnosis of Oligapta was made in order to accommodate O. kruidenieri n. sp. The presence of few testes, an unarmed cirrus, and simple clamp sclerites, the principal char- acters involved, are not considered at present to be sufficient to propose a new genus for this species. Discussion Views vary concerning the familial relationship of the Indocotylinae. Tri- pathi (1959) proposed the subfamily for his new genus and species, Indo- cotyle hemirhamphi, and placed it in the Discocotylidae primarily because of the presence of only 8 haptoral clamps. Tripathi (loc. cit.) did not mention the closely related genus Oligapta, which Unnithan (1957) had previously proposed in the Axininae of the Axinidae. Price (1962) recognized the In- docotylinae to include both Indocotyle and Oligapta but referred the subfamily to the Axinidae. Later, Yamaguti (1963) considered these genera unrelated to the Axinidae and replaced them in the Discocotylidae, while Young (1968) followed Price’s classification. Based on the comparative in- ternal morphology and the structure of the clamps, we conclude that the Indocotylinae belong in the Axinidae as originally proposed by Price (loc. cit.). Acknowledgments We wish to thank Dr. Donn E. Rosen and Mrs. Norma Feinberg, Amer- ican Museum of Natural History, for identifying Thyrinops pachylepis (Giin- ther). VOLUME 92, NUMBER 4 723 Literature Cited Dillon, W. A., and W. J. Hargis, Jr. 1965. Monogenetic trematodes from the southern Pacific Ocean. 2. Polyopisthocotyleids from New Zealand fishes: The families Discocotylidae, Microcotylidae, Axinidae, and Gastrocotylidae.—Jn Biology of the Antarctic Seas, II, Antarctic Res. Ser. 5, Am. Geophys. Union, p. 251-280. Kritsky, D. C., F. M. Bilqees, and P. D. Leiby. 1972. Studies on Monogenea of Pakistan. I. Pseudochauhanea elongatus sp. n. (Gastrocotylidae: Gastrocotylinae) from the gills of Labeo rohita (Ham.).—Proc. Helm. Soc. Wash. 39:231—233. Price, E. W. 1962. North American monogenetic trematodes. X. The family Axinidae.—Proc. Helm. Soc. Wash. 29:1-18. Tripathi, Y. P. 1959. Monogenetic trematodes from fishes of India.—Indian J. Helm. 9:1-149. Unnithan, R. V. 1957. On the functional morphology of a new fauna of Monogenea on fishes from Trivandrum and environs. Part 1. Axinidae fam. nov.—Bull. Cent. Res. Inst., Univ. Kerala, Ser. C 5:27-122. Yamaguti, S. 1963. Systema helminthum. IV. Monogenea and Aspidocotylea.—Interscience Pub., N.Y. Young, P. C. 1968. Oligapta manteri sp. nov. (Axinidae: Monogenoidea) and Pseudothora- cocotyle scomberomori sp. nov. (Gastrocotylidae: Monogenoidea) from South Pacific fishes.—Jour. Helminthol. 42:411-420. (JWC) Department.of Zoology, Washington State University, Pullman, Washington 99163; (DCK) Department of Allied Health Professions, Idaho State University, Pocatello, Idaho 83209; (RJK) Department of Biology, Idaho State University, Pocatello, Idaho 83209. PROC. BIOL. SOC. WASH. 92(4), 1979, pp. 724-742 BATHYPONTIA SARS (COPEPODA: CALANOIDA): EIGHT SPECIES, TWO NEW, FROM THE CARIBBEAN SEA AND GULF OF MEXICO Georgiana B. Deevey Abstract.—Eight species of Bathypontia, including two new species, are now known from the Caribbean Sea; six of these species have been recorded from the Gulf of Mexico. The two new species, B. michelae and B. unis- pina, are described, the distinctive characters of the eight species are dis- cussed and figured, and a key to the females of the genus is included. Eight species of Bathypontia, including 2 new species, have been iden- tified from samples collected by Dr. Harding Michel on 3 cruises of the PILLSBURY in the Caribbean Sea and Gulf of Mexico. Ten valid species have been previously described: B. elongata Sars 1905, B. minor (Wolfen- den) 1906, B. sarsi Grice and Hulsemann 1965 (=B. minor:Sars 1907), B. spinifera Scott 1909, B. elegans Sars 1920, B. longiseta Brodsky 1950, B. similis Tanaka 1965, B. longicornis Tanaka 1965, B. regalis Grice and Hul- semann 1967, and B. intermedia Deevey 1973. Of these only the females are known for B. longiseta and B. intermedia, and only the males for B. longicornis and B. regalis. Four of these species have been recorded from these waters: B. minor (Wolfenden) and B. similis Tanaka by Park (1970) and B. elongata and B. sarsi (as B. minor Sars) by Owre and Foyo (1964, 1967, 1972). Other species now known from the Caribbean Sea are B. ele- gans, B. spinifera, and 2 new species of which only the females were col- lected. Six of these species are also known to occur in the Gulf of Mexico: B. elongata, B. sarsi, B. spinifera, B. similis, B. minor, and B. michelae n. sp. The species most frequently noted was B. similis, with records of 25 females but no males. Ten specimens, including 3 males and 5 females, of B. elongata were taken; 5 males of B. spinifera; one female, one male, and one juvenile B. elegans; 2 females and one male B, minor; and a female, an immature female, and a male B. sarsi—as well as a female of one new species and 2 females of the other. I am deeply indebted to Dr. Harding B. Owre Michel for the privilege of examining some of her samples from the Caribbean Sea and Gulf of Mexico. This work was partially supported by grant GA-36512 from the National Science Foundation. The diagnosis of the genus remains as previously given (Deevey, 1973), VOLUME 92, NUMBER 4 F25 but the key to the species of females must be modified to include the new species. Also, although Sars described the sarsi female as 2.6 mm long and Scott the spinifera female as 3.3 mm long, it is now apparent that these species may be similar in size so size cannot be used as a key character to differentiate them. ~] oo | Key to the Females of Bathypontia Me Mcaitine teimales,OVER-4 Mimi) LONE. “FE. Fale. of yo 'cce cb D¥ co Diener anos Z Matunestemales-less‘tham4 mm long ...... 5. .6..'becc cd cae de eee 3 . Last segment of PS at least twice as long as preceding segment, genital segment almost as long as 3 following segments........... . 5 SEES se Se I SPA Seca cer ae ae Pe B. elegans Sars Last segment of PS little longer than preceding segment, genital seg- ment slightly longer than following segment ......... B. elongata Sars Mh SeASVIMMetriCal IMVGOTSAMWIEW «3. os<4e< ues oaow arse oe ee oe se 4 BhsrsvimmMethacal ta GOTSAl VEEW =2te sc ince when Sule 6 os be Ded kd OMoas « 5 . Left side of ThS longer than right, apical spine of PS as long as or longer than leg, no outer edge spines leg 1...... B. longiseta Brodsky Right side of Th5 longer than left, apical spine of P5 twice as long as distal segment, outer edge spines on exopod segment 3 of leg 1 5 2 SERRE cols Cacce S ae eee Oe sea B. similis Tanaka - P5awith smgle apical spine; no acceSSOry Spine ...:.....0 0.2 c ee. 6 P5 with apical spine and small accessory spine .................08. q . Apical spine ca. 4 times as long as distal segment of P5...B. minor (Wolfenden) Apical spine the same length as distal segment of PS ............. MN re sak oaks) a cc eee uae sis NS nore one: oa. ok Pee eee B. unispina n. sp. . P5 asymmetrical, small 2nd spine asymmetrically placed on the 2 SS: oo SRI a 0 a See esc neenis O geanaeretn ene B. intermedia Deevey IES) SWART CLS (Lat ae A ese J ee 8 . ThS pointed in dorsal view, rounded in lateral view ................ 9 ThS pointed in dorsal and lateral view ... B. sarsi Grice & Hulsemann . Basal segment of 2nd maxilla with series of 1-1-2-3 bristles or setae, basal segment of maxilliped with 1 proximal, 1 middle and 3 distal SPiMesn(SCe HIgS. Il. Opes, 2. wean s oh neuconeert B. michelae n. sp. Basal segment of maxilla 2 with clusters of spinules and 1-2-3 bris- tles, basal segment of mxp with 1 middle and 2 distal spines (see IS eC) nal code 0g Sc’. LAM A giyats aintd bag acEye asain t B. spinifera Scott Bathypontia michelae, new species Figs. 1 and 2 Holotype.—Female, 3.25 mm long, collected on 3 November 1969. On 2 726 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Bs b Fig. 1. Bathypontia michelae, female: a and b, Dorsal and lateral views; c, Antenna 2, longer setae cut off; d, Mandibular palp and blade; e, Maxilla 1, longer setae cut off; f, Maxilliped. Scale on left margin for a and b; at upper right for c-f. Scales in mm. VOLUME 92, NUMBER 4 727 [ YU eG Yi —— SS — Fig. 2. Bathypontia michelae, female; a, Blade of mandible; b, Maxilla 2: c, Sth legs; d, Leg 1, setae cut off; e, Leg 2; f, Leg 3; g, Leg 4.(Scale at left for a, near center for b-g. Scales in mm. 728 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON slides to be deposited in the National Museum of Natural History, Smith- sonian Institution. Type-locality.—Pillsbury Cruise 6911, Station 14:19°02'’N, 65°38’W, northeast of Puerto Rico, depth of tow 785 m. See Fig. 3 for station loca- tions. Paratype.—Female, 3.14 mm long, collected on 16 April 1968 on Cruise P6803, Station 12: 24°21’N, 86°12’ W, in the Gulf of Mexico, from 1,900 m depth. On 2 slides to be deposited in the National Museum of Natural His- tory, Smithsonian Institution. Etymology.—This species is named for Dr. Harding Michel. Description of female.—Body oblong, head separated from Th1, Th4 and ThS separate, Th5 pointed in dorsal view, rounded laterally (Figs. la, b). Urosome ca. a third of cephalothorax length, genital segment swollen ven- trally, about the same length as the 2 succeeding segments, anal segment relatively long, caudal rami short. Well-developed rostrum rounded and bent downwards. Antenna | of 23 segments, extending to urosome past genital segment. Antenna 2 endopodite with total of 16 setae slightly longer than exopodite, which has a pronounced proximal protuberance as in most other species of Bathypontia (Fig. 1c). Blade of mandible with the Ist tooth large and removed, like a thumb, from the rest of the toothrow, which consists of serrations capped by 3 notched transparent teeth that are readily lost or broken, as is the large transparent spike-like cap on the Ist tooth (Figs. 1d, 2a). The 3 notched teeth are followed by 3 spike-like teeth, a cluster of setae or spines, and a large densely spinous bristle. This bristle is relatively small compared with those found in other species, such as B. minor, B. spinifera, and B. similis. Maxilla 1 (Fig. le) with 10 setae on the Ist inner lobe, 1 seta each on 2nd and 3rd inner lobes and basis, 10 setae on the exopodite, and 4 on the outer lobe. Maxilla 2 (Fig. 2b) strongly developed and, as in the other species, with 6 strong subequal curving setae plus 3 smaller ones on the distal portion; proximal portion with 1-1-2-3 bristles on rounded protu- berances, the 3 proximal with spinules at their bases. Maxilliped (Fig. 1f) weak, with 1 proximal, 1 middle, and 3 distal spines on basal segment. Leg 1 with a 1-jointed endopod, though traces of segmentation may be discerned, and with 3-jointed exopod with 2 slim outer edge spines on exopod segment 3, and with a long curving internal seta on B2. Legs 2—4 with 3-segmented exopods and endopods. B1 of legs 1-3 with a plumose seta and a bunch of hairs; B1 of leg 4 bare, but B2 has a small outer spine. B2 of leg 3 with a long strong outer spine (Figs. 2d—g). Leg 5 (Fig. 2c) 3-jointed, symmetrical, with a long distal spine and a small 2nd outer spine; the distal segment is 75-80% of the length of the distal spine. Remarks.—At first it was expected that these females would prove to be B. spinifera, since they were the right size and shape for that species and males were present. In fact, the B. spinifera female has not been docu- VOLUME 92, NUMBER 4 w29 “ain 70°W B. unispina * GULF B. michelae OF B. elongata * MEXICO B. spinifera W B.sarsi i) ° B. elegans @ 25°N 15° Fig. 3. Station locations in the Caribbean Sea and Gulf of Mexico for 6 species of Bathy- pontia. mented or recorded since it was described by Scott in 1909, although males have been recorded from the Sargasso Sea (Deevey, 1973) and from near the Cape Verde Islands (Roe, 1975). Also the 5th legs are similar to those described by Scott, and judging from his figure (1909, Pl. III, fig. 16) the proportions of the leg segments and relative length of the distal spine agree with this description of B. michelae n. sp. However, examination of the appendages revealed many differences. The blade of the mandible differs from that of B. spinifera (Fig. 7e) and B. similis (Fig. 9h) and is of the same type as that found in B. elongata, B. elegans (Fig. 9f), and B. intermedia, in that the Ist tooth is removed some distance from the rest of the toothrow, and the densely setose bristle is relatively smaller than in the other species. Also the setation of the basal segment of the maxilla 2 and the maxilliped differs from that found in B. spinifera (Figs. 7f, g) and B. similis (Figs. 9f, i), and the endopod of leg 1 is 1-jointed, whereas it is 2-jointed in those species. B. michelae n. sp. appears to be most closely related to B. sarsi in the setation of the appendages, but the ThS is rounded in lateral view and the distal spine of leg 5 is somewhat longer with respect to the distal seg- ment. 730 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Bathypontia unispina, new species Figs. 4 and 5 Holotype.—Female, 3.65 mm total length, cephalothorax 2.85 mm to tip of Th5, urosome 0.95 mm long, collected on 1 November 1969. On 2 slides, to be deposited in the National Museum of Natural History, Smithsonian Institution. Type-locality.—Cruise P6911, Station 11: 17°30’N, 69°30’W, in the eastern Caribbean south of Hispaniola, from 3,602 m depth (see ne 3); bottom depth 4,191 m. Etymology.—The specific name refers to the single distal spine on the distal segment of the female 5th leg. Description of female.—Body oblong, head separated from Th1, Th4 and 5 separate, 5th thoracic segment somewhat pointed in dorsal view, rounded laterally (Figs. 4a, b). Antenna 1 of 23 segments, extending to urosome. Urosome 33-34% of cephalothorax length; genital segment swollen ven- trally, as long as 2 following segments together. Caudal rami shorter than anal segment; the 3 caudal setae were broken off. Rostrum prominent, sin- gle, rounded and bent downwards, slightly indented at tip (Fig. 4g). Antenna 2 endopodite with a total of 15 setae on the distal segments, slightly longer than exopodite (Fig. 4f). Mandible blade with Ist tooth removed from sev- eral following, its transparent cap missing, but probable outline indicated by broken lines in Figures 4d and e. Densely spinous bristle close to toothrow and relatively small. Mandible palp longer than blade with 5 setae on exop- odite and 9 long and | short setae on endopodite. Maxilla 1 (Fig. 4c) with 9 setae of varying lengths on Ist inner lobe, 2nd and 3rd inner lobes and basis each with 1 seta, 10 setae on exopodite, and 4 on outer lobe. Maxilla 2 strongly developed (Fig. Sa), distal part with 6 strong subequal setae plus 3 shorter ones, proximal part with setae in sequence of 1-1-2-3, the 4 most proximal setae with spinules at their bases. Maxilliped (Fig. 4h) weak, basal segment with setae in sequence of 1-1-3. Leg 1 (Fig. 5b) with 1-segmented endopod, 3-segmented exopod, 2 outer edge spines on exopod segment 3; B2 with a distal internal long curving seta. Legs 2—4 with 3-segmented branches. B1 of legs 1-3 with a plumose seta and a bunch of hairs on the inner edge (Figs. Sb—d). B1 of leg 4 without seta or hairs (Fig. 5e). Legs 2 and 4 with sharp external denticle on B2. Leg 3 B2 with long slightly curved external spine almost as long as exopod segment | (Fig. 5d). Exopod seg- ment 2 of leg 2 with a slightly longer outer edge spine than on segments 1 and 3. Plumose setae of legs 1—4 jointed. Leg 5 (Fig. Sf) 3-jointed, sym- metrical, basal segment slightly longer than 2nd segment, 3rd segment slight- ly more than twice as long as 2nd segment, with a single long distal spine as long as the 3rd segment. Remarks.—B. unispina n. sp. is similar in size to B. intermedia, but the VOLUME 92, NUMBER 4 731 Fig. 4. Bathypontia unispina, female: a and b, Dorsal and lateral views; c, Maxilla 1, longer setae cut off; d, Palp and blade of mandible; e, Mandibular blade; f, Antenna 2, most setae cut off; g, Rostrum; h, Maxilliped. Outer scale on right for a and b, other scale for c-h. Scales in mm. 732 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 5. Bathypontia unispina, female: a, Maxilla 2; b, Leg 1; c, Leg 2; d, Leg 3; e, Leg 4; f, 5th legs. Scale for a-f in mm. VOLUME 92, NUMBER 4 733 latter species has asymmetrical 5th legs. These 2 species are intermediate in size between the larger B. elegans, B. elongata, and B. longicornis and the other species which are all smaller. The 5th legs of B. unispina n. sp. differ from those of all other known Bathypontia females. In the other females there is a small 2nd outer distal spine, except in B. minor, in which the leg segments are shorter and the distal spine is four times as long as the distal segment. The mandibular blade of B. unispina n. sp. is of the same type as in B. elegans, B. elongata, B. intermedia and B. michelae n. sp. The setation of maxilla 1 is the same as in B. sarsi. The setation of the basal segments of the maxilla 2 and maxilliped agrees with that of B. sarsi and B. michelae n. sp. and differs from that of B. similis and B. spinifera. The endopod of leg 1 is 1-segmented as in B. sarsi, B. minor, B. michelae n. sp. and B. elegans. In the setation of the appendages this species is most closely related to B. sarsi and B. michelae n. sp. Bathypontia similis Tanaka Fig. 9g-i Bathypontia similis Tanaka, 1965:45, figs. 24la—k. Bathypontia similis.—Park, 1970:543, figs. 386—402. Bathypontia similis.—Deevey, 1973:365, figs. 3c, f, g, 4f. Material.—25 females, 2.75-3.0 mm long. The 17 station locations where the females were found are shown by the black stars in Figure 6; the region extended from 24°21'N in the Gulf of Mexico to 10°22’N off the coast of Central America, and from 61°30’W in the eastern Caribbean to 87°27’W off Belize. They were caught over a total depth range of 490-—3,500 m. The specimens recorded from the Sargasso Sea (Deevey, 1973) were taken from between 500 and 2,000 m-depths. The fe- male is readily identified by the asymmetry of the 5th thoracic segment, the right side being longer than the left. The proportions of the female 5th leg are similar to those of B. sarsi, but the long distal spine is twice as long as the distal segment, whereas in B. sarsi the distal segment is proportionately longer. The setation of the basal segments of the maxilla 2 and maxilliped is the same as in B. spinifera, as is the structure of the mandibular blade (see Figs. 9g—1). The endopod of leg | is also 2-jointed, as it is in B. spinifera, B. intermedia, B. regalis and B. elongata. It is curious that so many females were caught and no males. Tanaka’s (1965) description of the species was based on 4 females and 2 males. The only male documented since that time was taken in the Sargasso Sea (Deev- ey, 1973, Figs. 4f). Grice and Hulsemann (1967) listed B. similis from the Indian Ocean, but did not state the number or sex of the specimens. Distribution.—Pacific coast of Japan, 2°38’S in the Indian Ocean, 10°- 734 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON g5° 75° 65°w GULF OF ATLANTIC co met OCEAN 25°N CARIBBEAN 15° Oh? SOUTH AMERICA Fig. 6. Station locations for Bathypontia similis shown by black stars, for B. minor by black circles. 27°N in the Caribbean Sea and Gulf of Mexico, 32°10’N in the Sargasso Sea off Bermuda. Bathypontia spinifera Scott Fig. 7 Bathypontia spinifera Scott, 1909:183, pl. 3, figs. 1-16. ?Bathypontia sarsi.—Wheeler, 1970:12, figs. 77-90. Bathypontia spinifera.—Deevey, 1973:367, figs. 4a, b, g. Bathypontia spinifera.—Roe, 1975:364, figs. 3la—d. Material.—S5 males, 2.6—2.92 mm long. Males 2.8—2.92 mm long were caught over a depth range of 750—3,000 m in the western Caribbean and the Florida Current north of Cuba (see the small open stars in Fig. 3) at: 20°04’N, 79°38.4’W; 16°40’N, 87°W; 18°10’N, 87°W; and 23°30'N, 83°30’W. A 2.6 mm male was collected at 27°N, 86°W in the Gulf of Mexico in a 1,000-1,500 m haul. The 4 males from the Sargasso Sea were taken from 500-2,000 m-depths. Roe’s male was collected in a VOLUME 92, NUMBER 4 735 b Fig. 7. Bathypontia spinifera, male: a and b, Lateral and dorsal views; c, Maxilla 1, setae cut off; d, Antenna 2, setae cut off; e, Blade of mandible; f, Maxilliped; g, Basal segments of maxilla 2; h, Leg 1; i, Leg 2; k, Leg 3; m, Leg 4; n, 5th legs. Scale at bottom right for a and b; at bottom left for i-n; on center right margin for e; upper center for c, d, f-h. Scales in mm. 736 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON tow from 610-700 m off the Cape Verde Islands. The type-specimens were caught in a 1,000—0 m haul. Without giving any further data, Grice and Hulsemann (1967) listed B. spinifera from 8 hauls in the Indian Ocean all from between 1,000 and 2,000 m. As noted, the setation of the appendages is similar in B. similis and B. spinifera, as is the shape of the mandibular blade and the size of the densely spinous bristle (see Figs. 7c—g and 9g—i). And it is curious that so many females of B. similis have been recorded and only | male, whereas only male B. spinifera have been documented since the original description. As described and figured by Scott the female B. spinifera differs from female similis in having a symmetrical ThS, and in the relative lengths of the long distal spine and the distal segment of the 5th legs. In B. similis the distal Spine is twice the length of the distal segment, whereas, as drawn by Scott (1909, Pl. 3, Fig. 16), in B. spinifera the distal segment is longer, two-thirds the length of the distal spine. The males of both species have a symmetrical ThS and the Sth legs are fairly similar (see Figs. 7n and Deevey, 1973, figs. 4f and g). The long spine of the right exopod segment 2 of leg 2 (Fig. 71) of B. spinifera males is not as long on this segment in the B. similis male. Distribution.—1°4.5'S, 127°52.6’E in Indonesian Seas, 5°48’N—27°31’S in the Indian Ocean, 16°20’—27°N in the Caribbean Sea and Gulf of Mexico, 32°N in the Sargasso Sea and 18°N, 25°W in the eastern Atlantic. Bathypontia minor (Wolfenden) Fig. 9j, k, m Isocalanus minor Wolfenden, 1906:36, pl. 12, figs. 1-5; 1911:349, pl. 38, figs. 6-8, textfig. 78a, b. Bathypontia minor.—Grice and Hulsemann, 1965:249, figs. 19(e)—(q). Bathypontia minor.—Park, 1970:541, figs. 372-385. Bathypontia minor.—Deevey, 1973:365, figs. 3a, b, d, e, h, 1. Material.—2 females, 2.65 and 2.8 mm long, and 1 male 2.4 mm long. The females were caught at 18°15’N, 79°20’W and at 20°04’N, 79°38.4’W and the male at 20°02’N, 82°48’W, in the western Caribbean around the Cayman Islands south of Cuba, over a depth range of 890—1,950 m (see the closed circles in Fig. 6). Other recorded depths are 980—1,900 m in the Caribbean Sea and Gulf of Mexico (Park, 1970), 580-940 m in the Canary Island region (Roe, 1972), 0-1,500 m in the Sargasso Sea (Deevey, 1973), 1,000 m in the North Atlantic (Grice and Hulsemann, 1965), and 0—1,500 m (Wolfenden, 1906). This species has not been recorded from as great depths as most of the other species. B. minor is distinguished by the long distal spines on the male and female Sth legs, both sexes lacking a small second spine. The setation of the ap- pendages is also distinctive, especially that of the basal segments of the VOLUME 92, NUMBER 4 737 maxilla 2 and maxilliped, which have fewer setae or bristles than in other species, as well as the shape of the mandibular blade with its exceptionally large spinous bristle (Figs. 9j, k, m). Distribution.—Ca. 16°-32° N in the Atlantic, 18°-22°N in the Caribbean Sea and Gulf of Mexico. Wolfenden’s (1906) specimen was caught west of the Cape Verde Islands. B. minor has been recorded only from the North Atlantic and adjacent waters. Bathypontia sarsi Grice and Hulsemann 1965 Fig. 8 Bathypontia minor Sars, 1907:27; 1924: pl. 127, figs. 12-18; 1925:360. Bathypontia sarsi Grice and Hulsemann, 1965:249. Bathypontia sarsi.—Roe, 1975:361, figs. 30a-i. Material.—Female 2.9 mm long, juvenile female 2.3 mm long, male 2.8 mm long. The female was caught at 17°18'N, 85°27'W at 480 m-depths, the male at 16°09.5'’N, 76°12.2'W from 770 m, and the immature female at 27°30'N, 87°10'W in a vertical ‘tow (see Fig. 3). The male and female appendages are shown in Fig. 8, except for legs 1-4, which are as in the other species. The mandibular blade is distinctive (Figs. 8e, k) and has a relatively small spi- nous bristle as in B. michelae n. sp., B. unispina n. sp., B. intermedia, and B. elongata. The setation of the basal segment of maxilla 2 (Fig. 8g) is the same as in B. elegans, B. michelae n. sp., B. unispina n. sp., and B. elongata. The setation of the basal segment of the maxilliped (Fig. 8f) is as in B. michelae n. sp. and B. unispina n. sp. The endopod of leg 1 is 1- jointed, as in B. minor, B. elegans, B. unispina n. sp., and B. michelae n. sp. The male Sth leg is distinctive and notably asymmetrical, the right leg being longer with a triangular distal segment, and is quite different from the 5th legs of other species. Sars (1924, Pl. 127, fig. 18) figured a longer distal spine on the left leg than was present on our specimen or Roe’s (1975, fig. 31s) males. These specimens agree in detail with those Roe described and figured that were caught near the Cape Verde Islands. Roe concluded his 2 females and 3 males were indeed B. sarsi after comparing them with a male and a female B. sarsi from Sars’s material that he borrowed from the Oceanographic Museum in Monaco. The toothrow of the mandibular blade of his and our specimens differs somewhat in shape from Sars’s (1924, Pl. 127, fig. 16) figure of the mandible blade of his female. Sars’s figure shows a blade more like that of B. similis and B. spinifera. As B. minor Sars, B. sarsi was previously recorded from these waters by Owre and Foyo (1964, 1967, 1972) and their photograph (1967, figs. 725) of the mandible blade also shows a large setose bristle and the shape of the blade more as in those 2 species. 738 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 8. Bathypontia sarsi: a, Lateral view of male; b, Dorsal view of female; c, Female antenna 2, setae cut off; d, Female maxilla 1, setae cut off; e, Female mandibular blade and palp, most setae cut off; f, Female maxilliped; g, Basal segments of female maxilla 2; h, Female Sth legs; i, Male 5th legs; k, Blade of female mandible. Scale on right margin for a and b; bottom left for k; lower center for c—i. Scales in mm. VOLUME 92, NUMBER 4 739 Wilson (1950) has also recorded this species from the Pacific near the Philippines and from the China Sea off Formosa; his figure of the female 5th legs (1950, Pl. 22, fig. 303) could as easily represent the Sth legs of the B. similis female. Other records of this curously rare and elusive species (Lysholm, Nordgaard and Wiborg, 1945; Grice and Hulsemann, 1967) have not been documented in any way. Distribution.—Sars’s specimens are believed to have been collected near the Azores, so the documented occurrence of B. sarsi is from 18° to ca. 36°N in the eastern Atlantic, and from 15°—27°30'N in the Caribbean Sea and Gulf of Mexico. Bathypontia elongata Sars Bathypontia elongata Sars, 1905:24; 1924: pl. 126, figs. 1-17; 1925:356. Wsocalanus major Wolfenden, 1906:37, pl. 12, figs. 6-11; 1911:347, pl. 38, figs. 3-5, textfig. 77. Bathypontia elongata.—Farran, 1908:87, pl. 9, figs. 16, 17. Bathypontia elongata.—Grice and Hulsemann, 1967:38, figs. 271-274. Bathypontia elongata.—Deevey, 1973:367, figs. 4c—e, 5a, b, d, g, h. Material.—S females, 5.2—5.6 mm long, 3 males 4.9—5.2 mm long, and an immature male and an immature female, both ca. 4 mm long. The 8 station locations where B. elongata was captured over a depth range of 485—1,700 m are indicated by the small black stars in Fig. 3. They ranged from 9°50’N off Central America and from 67°14’W in the eastern Caribbean to 28°12'’N, 88°37’ W in the Gulf of Mexico. Owre and Foyo (1972) have also reported this species from a depth of 435 m at 15°N, 64°W. This species is distinctive largely because of its size and possibly because it lives at shallower depths than most of the other species. It has been recorded from the Atlantic, Pacific, and Indian Oceans by a number of investigators (Farran, 1908; Lysholm, Nordgaard, and Wiborg, 1945; Jes- persen, 1940; Wilson, 1950; Vervoort, 1965; Grice and Hulsemann, 1967; Roe, 1972; Deevey, 1973). Distribution.—In the Atlantic from the Gulf of Guinea at 1°30'N to 63°38'N south of Iceland, and in the Caribbean Sea and Gulf of Mexico, and also from ca. 5°—23°S in the eastern Pacific, and 18°N-—12°S in the Indian Ocean. Bathypontia elegans Sars Fig. 9a—f Bathypontia elegans Sars, 1920:26; 1924: pl. 127, figs. 1-11; 1925:358. Bathypontia elegans.—Deevey, 1973:369, figs. 5c, e, f, i-k. 740 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON I BB A) a ee a BN Ger Fig. 9. Bathypontia elegans, a-f: a, Dorsal view of male with left antenna 1; b, Lateral view of male with right antenna 1; c, Basal segment of male maxilliped; d, Male Sth legs; e, Basal segments of male maxilla 2; f, Mandible blade of immature female. Bathypontia similis female, g-i: g, Basal segment of maxilliped; h, Mandible blade; i, Basal segments of female maxilla 2. Bathypontia minor, j-m: j, Blade of female mandible; k, Basal segments of female maxilla 2; m, Male maxilliped. Upper scale on left margin for a and b; at bottom left for d; on f for f; at top right for c-e, g, i, k, m. Scales in mm. VOLUME 92, NUMBER 4 741 Material.—Male 4.2 mm long, female 4.2 mm long, and an immature female 3.4 mm long. The male was caught at 19°15’N, 67°15’W, the female at 21°08’N, 74°42'W, and the immature female at 20°28’N, 72°30'W, over a depth range of 1,590-2,232 m (see the closed circles in Fig. 3). The females from the Sargasso Sea (Deevey, 1973) were found between 1,000 and 2,000 m-depths. The only other record of this species, since it was described by Sars, is Grice and Hulsemann’s (1967) undocumented report of it as B. major from the Indian Ocean. B. elegans is almost as large as B. elongata, but is easily distinguished from that species by the shape of the rostrum, the Sth legs, and the pro- portions of the urosome. In the B. elegans female the distal segment of the 5th legs is much longer in proportion to the distal spine than in the B. elongata female. The B. elegans male Sth legs (Fig. 9d) have exceptionally long distal spines; only the B. minor male has such long distal spines, but that species is much smaller and lacks the 2nd outer small spine on both 5th legs. The median seta on the male left caudal ramus is exceptionally long (Fig. 9a). The mandibular blade (Fig. 9f) is of the same type as in B. elongata, B. intermedia, B. unispina n. sp., and B. michelae n. sp., and has the Ist tooth removed from the rest of the toothrow and a small spinous bristle. The setation of the basal segments of maxilla 2 (Fig. 9e) is the same as in B. elongata, B. sarsi, B. michelae n. sp., and B. unispina n. sp. The basal segment of the maxilliped has 2-2-3 spines (Fig. 9c), more than in any other known species. The male leg 2 is asymmetrical, with a longer long spine on the right exopod segment 2 than on the left exopod segment 2, although it is not as exceptionally long as in the B. spinifera male (Fig. 71). Distribution.—19°—32°N in the western Atlantic, 33°—38°N in the eastern Atlantic, and 22°11’S in the Indian Ocean. Literature Cited Brodsky, K. A. 1950. Calanoida of the Far Eastern Seas and Polar Basin of the USSR (In Russian; Translated 1967). Keys to the fauna of the USSR, No. 35.—Zool. Inst. Acad. Sci. USSR, Moscow-Leningrad, 440 pp. Deevey, G. B. 1973. Bathypontia (Copepoda: Calanoida): Six species, one new, from the Sargasso Sea.—Proc. Biol. Soc. Wash. 86:357—372. Farran, G. P. 1908. Second report on the Copepoda of the Irish Atlantic Slope.—Fisheries, Ireland, Sci. Invest., 1906, II. (1908), 104 pp., 10 pl. Grice, G. D., and Kuni Hulsemann. 1965. Abundance, vertical distribution and taxonomy of calanoid copepods at selected stations in the northeast Atlantic.—Jour. Zool. 146 (2):213-262. , and . 1967. Bathypelagic calanoid copepods of the western Indian Ocean.— Proc. U.S. Nat. Mus. 122 (3583): 1-67. Jespersen, P. 1940. Non-parasitic Copepoda.—The Zoology of Iceland 3 (33):1—166. Lysholm, B., O. Nordgaard, and K. F. Wiborg. 1945. Copepoda from the ‘‘Michael Sars”’ 742 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON North Atlantic deep-sea expedition 1910.—**Michael Sars’’ North Atlantic Deep-Sea Exped. 1910. 5 (7):1-60. Owre, H. B., and M. Foyo. 1964. Report on a collection of Copepoda from the Caribbean Sea.—Bull. Mar. Sci. Gulf Carib. 14 (2):359-372. , and . 1967. Copepods of the Florida Current. Fauna Caribaea No. 1, Crustacea, Part 1: Copepoda.—Institute of Marine Science, University of Miami, p. 1—137, 900 figs. , and . 1972. Studies on Caribbean zooplankton. Description of the Program and results of the first cruise.—Bull. Mar. Sci. 22 (2):483-521. Park, T. 1970. Calanoid copepods from the Caribbean Sea and Gulf of Mexico. 2. New species and new records from plankton samples.—Bull. Mar. Sci. 20 (2):472—546. Roe, H. S. J. 1972. The vertical distributions and diurnal migrations of calanoid copepods collected on the SOND Cruise, 1965. I. The total population and general discussion.— Jour. Mar. Biol. Ass. U. K. 52:277-314. 1975. Some new and rare species of calanoid copepods from the northeastern At- lantic.—Bull. Brit. Mus. (Nat. Hist.) Zool. 28 (7):297-372. Sars, G. O. 1905. Liste préliminaire des Calanoidés recueillis pendant les campagnes de S. A. S. le Prince Albert de Monaco, avec diagnoses des genres et des espéces nouvelles. Part 1.—Bull. Mus. Océanogr. Monaco, No. 26:1-24. . 1907. Notes supplémentaires sur les Calanoidés de la Princesse-Alice.—Bull. |’Inst. Océanogr. Monaco, No. 101:1-—27. . 1920. Calanoidés recueillis pendant les campagnes de S. A. S. le Prince Albert de Monaco. (Nouveau supplément).—Bull. Inst. Océanogr. Monaco. No. 377:1-—20. . 1924, 1925. Copépodes particulicgrement bathypélagiques provenant des campagnes scientifiques du Prince Albert 1 de Monaco.—Résult. Camp. Scient. Prince Albert I, Fasc. 69, pls, 1-127 (1924), pp. 1-408 (1925). Scott, A. 1909. The Copepoda of the Siboga Expedition. Part I. Free-swimming, littoral and semi-parasitic Copepoda.—Siboga Exped., Monogr. No. 29a, 323 pp., 69 pls. Tanaka, O. 1965. The pelagic copepods of the Izu Region, Middle Japan. Systematic Account XIII. Parapontellidae, Acartiidae and Tortanidae.—Publ. Seto Mar. Biol. Lab. 12 (5):379-408. Vervoort, W. 1965. Pelagic Copepoda. Part II. Copepoda Calanoida of the families Phaennidae up to and including Acartiidae, containing the description of a new species of Aetidei- dae.—Atlantide Rep. No. 8:9-216. Wheeler, E. H., Jr. 1970. Atlantic deep-sea calanoid Copepoda.—Smithsonian Contrib. Zool., No. 55:1-31. Wilson, C. B. 1950. Copepods gathered by the United States Fisheries Steamer *‘Albatross”’ from 1887 to 1909, chiefly in the Pacific Ocean.—U. S. Nat. Mus. Bull. 100, 14 (4):1- 441, 36 pls. Wolfenden, R. N. 1906. Plankton Studies. Part 2. Copepoda.—Rebman Ltd., London, pp. 25—44, pls. 8-14. . 1911. Die Marinen Copepoden der Deutschen Siidpolar-Expedition 1901-1903. II. Die Pelagischen Copepoden der Westwinddrift und des Stidlichen Eismeers mit Beschrei- bung mehrerer neuer Arten aus dem Atlantischen Ozean.—Deutsche Siidpolar-Exped. 12 (Zool. 4): 182-380, pls. 22—40. Florida State Museum, University of Florida, Gainesville, Florida 32611. PROC. BIOL. SOC. WASH. 92(4), 1979, pp. 743-752 DESCRIPTION OF THE LARVA AND PUPA OF CYLORYGMUS LINEATOPUNCTATUS (COLEOPTERA: HYDROPHILIDAE: RYGMODINI) Paul J. Spangler Abstract.—The larva, pupa, habitat, and distribution of the Chilean water beetle, Cylorygmus lineatopunctatus d’ Orchymont, are described and illus- trated. The larva and pupa are interpolated into previously published keys to the immature stages of hydrophilid beetles. In late January and through February, 1978, I was able to participate in entomological fieldwork in Chile through the combined auspices of the J. I. Molina Institute of Studies and Publications of Chile and the Smithsonian Institution. During that cooperative project I concentrated my efforts on collecting aquatic Coleoptera, and I was pleasantly surprised to find adults and the larval and pupal stages of the rare rygmodine hydrophilid beetle, Cylorygmus lineatopunctatus d’Orchymont. As mentioned previously (Spangler, 1974), C. lineatopunctatus is the only representative of the tribe Rygmodini known from South America. All other taxa of the Rygmodini, ca 18 genera and ca 53 species, have been described from New Zealand, the subantarctic islands of New Zealand, or from Australia. Larvae of two rygmodine genera, Thomosis and Namostygnus, were described recently by Ordish (1974), therefore, the following description adds a third larval description to our knowledge of the Rygmodini. The description of the pupa of Cylorygmus given below is the first for any of the rygmodine genera. In addition to describing the larva and pupa of C. lineatopunctatus, I am taking this opportunity to summarize and comment on new information on the bionomics and distribution of the species. The monotypic genus Cylorygmus was described by d’Orchymont (1933) from a single specimen which he found among hydrophilids in the Hamburg Museum. The type-species of the genus, C. lineatopunctatus d’Orchymont, was a female collected in 1896. Since d’Orchymont’s description of the genus and species in 1933, the species was referred to in the literature only five times as follows. Blackwelder (1944) cited C. lineatopunctatus in his checklist of Latin American Coleoptera. Moroni (1973) included it in his list of water beetles known from Chile. Spangler (1974) designated a neotype for the type which was destroyed when the Hamburg Museum was damaged during World War II, illustrated the neotype and distinguishing characters, 744 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON y Ve Wo. ( ee Fig. 1. Cylorygmus lineatopunctatus d’Orchymont, larva, habitus view. VOLUME 92, NUMBER 4 745 and discussed what little was known regarding its bionomics. Camousseight and Moroni (1976) discussed the species briefly, stating that the type was in the Hamburg Museum, not realizing it had been destroyed, and gave locality-data for two specimens in the collections of the Museo Nacional de Historia Natural de Chile. Bachmann (1977) in a section on the Hydrophil- idae in a compilation of taxonomic references on the aquatic biota of austral South America, cited d’Orchymont’s article in which C. lineatopunctatus was described. The specimens referred to below are either in the U.S. National Museum of Natural History, Smithsonian Institution (USNM) or the Museo Nacional de Historia Natural de Chile (MNHN). Larva Body (Fig. 1) elongate, slightly compressed dorsoventrally; habitus dis- tinctive because of large tubercles on body. Total length 8.8 mm; greatest width of prothorax 1.0 mm. Color of integument light yellowish brown. Sclerotized head capsule, thoracic terga and sterna, tergal sclerites of ab- domen, spiracles, and legs yellowish brown. Integument covered with as- perities which are generally oriented transversely except around tubercles and spiracles (Fig. 11) where they are arranged concentrically. Head quadrangular (Fig. 3); 0.8 mm wide, 0.6 mm from labroclypeus to occipital foramen. Frontoclypeal suture feebly indicated. Frontal sutures widely separated at base of head, therefore, epicranial suture absent. Frons not sagittate (Fig. 3). Cervical sclerites absent. Lateral margins behind ocelli with several (6 to 9) long, stout, yellow setae. Ventral surface of head with numerous seta-bearing punctures laterally and irregular longitudinal row of 8 seta-bearing punctures on discal area on each side of midline; with 2 deep, approximate, posterior tentorial pits behind gula. Labroclypeus prominent (Fig. 5), almost symmetrical; with 1 narrow me- dial tooth between 2 broadly rectangular teeth; broad teeth notched apically; left broad tooth slightly shorter than right broad tooth; 4 stout setae present, 1 on medial side of each broad tooth and 1 in notch of each broad tooth. Anterolateral projections of epistoma shorter than teeth; each projection with 3 or 4 stout setae anteriorly. Ocular areas each with groups of 6 distinct ocelli arranged in an ellipse. Anterior 3 ocelli larger and close to each other; posterior 3 ocelli smaller, ventrolateral one smallest and separated from posteromedial two. Antenna (Fig. 8) short, cylindrical, and twice as long as stipes; first seg- ment longest, about one-third longer than penultimate segment; penultimate segment with a long slender apicomedial seta and a long moderately stout tubercle, tubercle very closely appressed to ultimate segment, therefore, 746 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON & =I ES i “Sy i Figs. 2-11. Cylorygmus lineatopunctatus d’Orchymont: 2, left mandible, dv; 3, head, dv; 4, right mandible, dv; 5, labroclypeus, dv; 6, maxilla, dv; 7, spiracular atrium, dv; 8, antenna, dv; 9, labium, dv; 10, spiracular tubercle, dv; 11, labium, vv. dv = dorsal view, vv = ventral view. VOLUME 92, NUMBER 4 747 difficult to see; ultimate segment small and slender, about one-half as long as penultimate segment. Mandibles (Figs. 2, 4) symmetrical, prominent, stout, sharply pointed apically; each mandible with 2 large, well-defined inner teeth and 1 large distal tooth. Maxilla (Fig. 6) with stipes stout but short, about as long as palpifer; bearing 2 stout apicomedial, 3 basomedial, and | apicolateral setae. Palpifer segmentlike; with short, slender, sclerotized appendage on apicomedial an- gle; appendage about one-tenth as long as palpifer and bearing 2 apical setae. Palpus tapering distally; first segment subequal to penultimate; penultimate segment bearing a small slender seta apicomedially; ultimate segment short- est, about two-thirds as long as penultimate segment. Labium (Figs. 9, 10) extending to midlength of palpifer. Penultimate seg- ment of palpus elongate, cylindrical; ultimate segment more slender and slightly shorter than penultimate segment, and bearing an apical seta. Ligula distinct, apparently segmented, subequal in length to labial palpus. Mentum pentagonal; dorsal surface with several short setae laterally and 2 short setae near base of ligula; ventral surface glabrous except for a long slender seta on each side of midline in discal area. Submentum arcuate laterally; narrower than mentum widening posteriorly; dorsal surface with 5 or 6 very coarse spines along anterior edge and 4 irregular, longitudinal rows of spines, 1 row on each lateral margin and a pair paralleling midline; ventral surface glabrous except for a few apicolateral setae. Gula pentagonal, rounded posteromedially. Prothorax with sides nearly straight but diverging slightly posteriorly; posterolateral angles broadly rounded; anterior margin with 6 long robust setae, 3 equally spaced on each side of sagittal line; lateral margins each with 3 similar equally spaced robust setae; each lateral margin with a round- ed tumescence at midlength; sagittal line present. Prosternal sclerite large, subrectangular; with slightly notched area posteromedially suggesting an incomplete sagittal line; sides of sclerite narrow and extending around sides and into cervical region, apparently uniting on midline. Mesothorax wider than prothorax and about two-thirds as long (measured on midline) as prothorax; with 4 narrow, transverse, anterior sclerites and 2 large subrectangular mesotergal sclerites with anterolateral angles nar- rowed and extended anteriorly; each lateral margin with a prominent spi- racular tubercle which is preceded by a moderate fleshy tubercle and fol- lowed by a very large inverted coneshaped setiferous tubercle; sagittal line present. Metathorax slightly wider than and as long as mesothorax; anterior meta- tergal sclerites transverse, larger than posterior sclerites, irregularly rect- angular; posterior sclerites transverse, small, narrow; sagittal line present; 748 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON each lateral margin with 3 setigerous tubercles, 2 smaller, pleural tubercles, and | very large lateromedial tubercle. Legs four segmented; procoxae large, separated by a distance equal to length of profemur; trochanter about two-thirds as long as femur (viewed ventrally); femur slightly longer than tibiotarsus; tarsal claw single, ventrally without setae. Abdomen of 8 distinct segments, segments 9 and 10 reduced; terga similar to each other and separated by intersegmental membrane; segments 1 through 7 each with a pair of small, oval, well-sclerotized tergites. True segmentation obscured by additional transverse folds on segments; segmen- tal folds continued onto sternum. Each segment with 2 folds; anterior fold with 8 tubercles, 2 pleural, 2 lateral, 2 pre-spiracular, and 1 on each side of midline; posterior fold with 4 small tubercles, 2 lateral and 2 sublateral. All tubercles densely covered with asperities. A large spiracular tubercle present laterally on segments | through 7, each spiracle located laterally and slightly behind a large fleshy setiferous tubercle. Epipleurites and hypopleurites moderately lobed. Eighth tergum represented by superior valve of stigmatic atrium which bears a single large sclerite (Fig. 7), beneath which lies the eighth pair of abdominal spiracles. Ninth tergum with a single, small, medial, rectangular sclerite. The larva of Cylorygmus lineatopunctatus runs to couplet 23 in Ber- trand’s (1972) key to hydrophilid larvae, but it may be distinguished im- mediately from the other genera keying to that couplet because it has 3 apicomedial teeth on the labroclypeus instead of the 4 or more reported for the other genera which run to that couplet. Furthermore, the conspicuous tubercles on the abdomen provide the larva of Cylorygmus with a distinctive habitus which makes it easy to recognize. Pupa Total length 5.0 mm, greatest width 2.7 mm. Color white except styli and cerci yellowish brown. Glabrous except for styli described below. Head with 2 supraorbital styli above each eye. Pronotum with 24 styli as follows: 3 on each anterolateral angle, 2 on each side of midline on anterior margin, 3 on each posterolateral angle, 2 on each side of median line at posterior margin and 2 on each side of midline on disc of pronotum. Mesonotum with 2 styli, 1 on each side of scutellum. Meta- notum with 2 styli, 1 on each side of midline. Abdomen with 4 styli on first segment. Segments 2 through 7 each with a pair of pleural and 4 tergal (total 6) styli arranged as follows: 1 pleural stylus lateral to each spiracle, 1 stylus behind each abdominal spiracle, and I stylus between each spiracle and midline. Segment 8 with 2 styli on pos- VOLUME 92, NUMBER 4 749 Figs. 12-13. Cylorygmus lineatopunctatus d’Orchymont, pupa; 12, dorsal view; 13, ven- tral view. terior margin, 1 on each side of midline. Segment 9 with 2 cerci slightly shorter than basal width of segment 9; each cercus terminating in a mod- erately incurved apex. Abdominal segments | through 8 each with a pair of spiracles; those on first and eighth segments very reduced. Antennae and femora extending outward at right angles from body axis. Maxillary palpi extending posteriorly almost parallel to body axis. Tibiae of all legs folded back against their respective femora. Tarsi turned backward and parallel with body axis. The very large pseudobasal segment of the maxillary palpus, the carinae on the hind tibiae visible through the pupal integument, and the partially developed parameres and median lobe of the male genitalia visible at the apex of the abdomen identify the pupa described above as a male. The pupa of Cylorygmus lineatopunctatus keys to couplet 6 in Bertrand’s (1972) key to hydrophilid pupae (i.e., to Cymbiodyta Bedel, Enochrus Zaitz- ev, and .Helochares Mulsant). Because the pupae of these genera are so 750 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON similar, no key has been prepared previously to separate them. Further- more, the three genera each include numerous species, and in most in- stances the pupa of only one species of each genus is known; so the key presented below must be considered provisional. 6a. Middle and hind legs with only four tarsal segments visible; U.S., Mexico;:and, Burope 40 «G05 3 os ale es ee eee Cymbiodyta — Middle and hind legs with five tarsal segments visible ........... 6b 6b. Basal segment of hind tarsus short but subequal to second segment; maxillary palpi short, robust; Chile ................... Cylorygmus — Basal segment of hind tarsus short, much shorter than second seg- ment; maxillary palpi elongate, slender; cosmopolitan ............ 6c 6c. Pseudobasal segment of maxillary palpus with concavity toward the front; mesosternum rarely protuberant or laminate .. Helochares — Pseudobasal segment of maxillary palpus with concavity toward the rear; mesosternum always protuberant or laminate .... Enochrus Habitat The single larva of C. lineatopunctatus described above was collected from debris removed from a logjam. The 2 pupae and 9 adults found with the pupae were collected from loam caught up in a smaller pile of small logs and driftwood, Both habitats were in midstream. The adults and pupae were found in compacted loam among roots of a weed which held the loam in a matlike fashion. The Rio Anticura flows through rocks of volcanic origin. Colorimetric water testing methods indicated a pH of 5 and 0 grains of hardness for the river. During the approximately 2 weeks spent collecting in the area the daytime air temperature varied from 54°F to 71°F. The water temperature in the river varied from 44°F to 48°F. In my previous discussion of C. lineatopunctatus (Spangler, 1974), I re- ported that the adults collected near the beach at Isla Desolacion by Dr. O. S. Flint, Jr., were found as if hibernating in small spaces between roots of mosses and grasses and the rocks he overturned as he searched for insects. These adults were found on 1 and 5 October 1969 which would be early summer at Isla Desolacion and in the Province of Magalianes, Chile. The occurrence of adults alongside rocks as mentioned above and early in the summer suggests that C. lineatopunctatus may overwinter in the adult stage. The absence of eggs and presence of a single larva, 2 pupae, and the 9 adults collected in Osorno Province in mid February, at the end of summer in the area, although sketchy evidence, also suggests that the species may overwinter in the adult stage. The occurrence of adults of C. lineatopunctatus among roots of mosses and grasses on the beach at Isla Desolacion agrees reasonably well with the VOLUME 92, NUMBER 4 IS SINUSOIDAL PROJECTION oe tena Fig. 14. Cylorygmus lineatopunctatus d’Orchymont, known distribution. type of habitats reported by Ordish (1974) for three subantarctic rygmodine species, Namostygnus flemingi Ordish, Namostygnus pictus (Kirsch), and Thomosis guanicola Brown. Adults of these species were collected in ter- restrial habitats in leaf mold or litter, among plant roots, among sea lion excreta, in seabird nests, and under stones in a penguin colony. Ordish further reported larvae of T. guanicola from guano and small plants in penguin colonies. Distribution When I published my previous account (Spangler, 1974) of C. lineato- punctatus, 1 had seen only the 12 adult specimens collected by Dr. O. S. Flint, Jr., in Magallanes Province in southern Chile. Since then, I have seen 11 adults, 2 pupae, and 1 larva from two additional localities which consid- erably increases the known north-south distribution (Fig. 14). The speci- mens examined (26) are all from Chile as follows. CHILE: Magallanes Prov- ince: Fiordo Peel, 1 Oct. 1969, O. S. Flint, Jr., 7 6d, 1 2 (neotype-series) (USNM); Isla Desolacion, Puerto Churruca, 5 Oct. 1969, O. S. Flint, Jr., fae PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 266, 2 22 (paratypes) (USNM). Osorno Province: Puyehue National Park, Anticura, in Rio Anticura, 3 Feb. 1978, P. J. Spangler, 1 larva (USNM); same locality, 12 Feb. 1978, P. J. Spangler, 5 66,4 2°, 2 pupae (USNM). Valparaiso Province: Quillota, Feb. 1897, 1 6 (MNHN); Quillota, 1 6 (MNHN). Moroni (1973) also reported C. lineatopunctatus from San- tiago Province at Lo Aguila. Acknowledgments I am very grateful to Luis E. Pena G. who organized the cooperative fieldwork between the J. I. Molina Institute of Studies and Publications of Chile and the Smithsonian Institution during which time the immature stages of C. lineatopunctatus were collected. For financial support for this coop- erative fieldwork I tender my thanks to the administrators of the J. I. Molina Institute and the Smithsonian Institution’s Fluid Research Fund. I also thank Mr. Michael Druckenbrod, Smithsonian Institution staff artist, for preparing the illustrations included in this article. Literature Cited Bachmann, A. 1977. Hydrophilidae. In: Biota Acuatica de Sudamerica Austral, pages 231- 237.—San Diego: San Diego State University. Bertrand, H. P. I. 1972. Larves et Nymphes des Coléoptéres Aquatiques du Globe. 804 pp., 561 figures.—Abbeville, France F. Paillart. Blackwelder, R. E. 1944. Checklist of the Coleopterous Insects of Mexico, Central America, the West Indies, and South America. Part 1.—Bull. U.S. National Museum 185:1—188. Camousseight, A., and J. Moroni. 1976. Los tipos de Coleopteros acuaticos depositados en la coleccion del Museo Nacional de Historia Natural de Chile (MNHN).—Noticiario Men- sual, 21(242—243):3-6. Moroni, J. B. 1973. Elenco sistematico, sinonimico y distribucion de Coleopteros acuaticos Chilenos.—Rev. Chilena Entomol. 7:193—200. d’Orchymont, A. 1933. Contribution a l’etude des Palpicornia. VIII.—Bull. Ann. Soc. Ent. Belgique 73:271-313. Ordish, R. G. 1974. Arthropoda of the Subantarctic Islands of New Zealand (3)* Coleoptera: Hydrophilidae.—Journ. Roy. Soc. New Zealand 4(3):307-314, 4 figures. Spangler, P. J. 1974. The Rediscovery of Cylorygmus lineatopunctatus (Coleoptera: Hydro- philidae: Sphaeridiinae: Rygmodini).—Journ. Kansas Entomol. Soc. 47(2):244—248, 7 figures. Smithsonian Institution, Washington, D.C. 20560. PROC. BIOL. SOC. WASH. 92(4), 1979, pp. 753-761 A NEW GENUS OF MADICOLOUS BEETLES FROM ECUADOR (COLEOPTERA: HYDROPHILIDAE: HY DROBIINAE) Paul J. Spangler Abstract.—A new hydrobiine hydrophilid water beetle from Ecuador, Dieroxenus cremnobates, new genus, new species, is described, illustrated, and interpolated into previously published keys to related genera. Notes on the biotope and the genera of hydrophilids with which the new taxon was associated in its madicolous habitat are included. During a search in madicolous habitats for aquatic beetles belonging to the genus Oocyclus, specimens of an unrecognized genus of hydrophilid beetles were encountered and collected on several occasions. An exami- nation of these beetles and comparisons with described genera indicate that they represent a new genus belonging to the subfamily Hydrobiinae, the tribe Hydrobiini, and the subtribe Hydrobiae. This new genus, presently known only from the Andean Cordillera of Ecuador, is described below. Hydrobiini Dieroxenus, new genus Head with clypeus expanded and shelflike in front of eyes. Clypeus strongly emarginate anteromedially but without membranous preclypeal area. Eyes, viewed from above, moderately transverse; feebly reniform; widely separated. Antenna 8 segmented; 2 basal, 2 intermediate, | cupule, and 3 club segments. Maxillary palpus robust; short, about as long as an- tennae; 4 segmented; basal segment very short, second (pseudobasal) seg- ment straight and more robust than subsequent segments; second and third segments about equal in length; fourth segment slightly longer than third segment. Mentum strongly marginate-foveate apicomedially. Prosternum not carinate. Mesosternum with a fine, very low longitudinal carina medially on posterior half and extending between mid-coxae. Front, middle, and hind femora densely pubescent on basal three-fourths, apices glabrous. Metatro- chanters not elongated. Metatibiae straight. Tarsal formula 5-5-5. Pronotum with posterolateral angles rounded. Elytron with sutural stria extending from apex to about basal fourth. Epipleura sharply declivous along entire length, not horizontal. Type-species of the genus.—Dieroxenus cremnobates, new species. Gen- der: masculine. 754 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Etymology.—Dieroxenus from dieros, G.—wet, plus xenos, G.—stranger; i.e., wet stranger, because the beetles were unknown and they were living on seeps or adjacent to waterfalls under a thin film of water trickling over the rock surfaces. The new genus described above does not fit easily into either of the two subtribes in d’Orchymont’s (1942) study of the tribe Hydrobiini. The new genus superficially resembles a convex species of Enochrus (subtribe Hel- ocharae) but close examination shows the following differences; the char- acteristics for Dieroxenus given first followed by those for Enochrus: (1) antenna 8 segmented vs. 9 segmented; (2) maxillary palpus robust, as long as antenna vs. usually slender and longer than antenna; (3) last segment of maxillary palpus articulated toward mouth vs. articulated outward away from mouth; (4) second segment of maxillary palpus straight vs. convex anteriorly; (5) preclypeus absent vs. present; (6) mentum strongly emargin- ate-foveate apicomedially vs. not emarginate-foveate; (7) mesosternum fee- bly longitudinally carinate vs. longitudinally laminate. Therefore, because the majority of its characteristics show greater affinities to the subtribe Hydrobiiae rather than to the Helocharae, I have assigned Dieroxenus to the Hydrobiiae. In d’Orchymont’s (1942) key to the genera belonging to the Hydrobiiae, Dieroxenus keys to the first rubric in couplet 13 because the prosternum is non-carinate. However, the mesosternum has a fine, low, longitudinal carina instead of being non-carinate and the metafemur is pubescent on the basal three-fourths instead of only basally or only along the foremost edge. The new genus described above may be distinguished from the others in the sub- tribe by interpolating the following in place of d’?Orchymont’s couplet 13. 13... “Prosternum. non-canmates 0). 2 Pe a ee eee 13a — Prosternum carinate longitudinally ........... Paracymus Thomson 13a. Mesosternum non-carinate longitudinally or with no longitudinal median carina behind the anterior transverse carina ............ 14 13b. Mesosternum with a fine very low, longitudinal carina between mid coxae and extending over posterior half of mesosternum (Fig. A eget i SuSe, en enc osen tae” eGR TOE ee Dieroxenus Spangler Dieroxenus cremnobates, new species Figs. 1-8 Holotype male.—Length 3.4 mm, greatest width 1.9 mm at midlength. Color of head piceous except margins in front of eyes dark reddish brown. Pronotum piceous discally and dark reddish brown laterally and along an- terior margin. Elytra very dark reddish brown, sides lighter reddish brown. Venter piceous except maxillary and labial palpi, and basal 5 segments of antennae yellowish brown; antennal club darker brown; legs reddish brown. VOLUME 92, NUMBER 4 755 5 if 2 x a See ty Fig. 1. Dieroxenus cremnobates, n. gen., n. sp., habitus view. Head finely, moderately densely punctate, punctures separated by one and one-half to two times their width except a few very coarse punctures at anteromedial corners of eyes and several at anterolateral corners of clyp- eus. Clypeus in front of eyes (Fig. 2) broadly expanded, shelflike; arcuately 756 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON emarginate anteromedially; without preclypeus. Labrum more finely but as densely punctate as head; feebly emarginate medially. Eyes separated by about 5 times their width. Ventral surface of head finely alutaceous behind eyes on gular region. Mentum deeply emarginate and deeply foveate api- comedially (Fig. 3) and finely, very sparsely punctate. Antenna 8 segmented (Fig. 2); 2 basals, 2 intermediates, 1 cupule, and 3 club segments. Maxillary palpus 4 segmented, about as long as antenna; basal segment very short; second (pseudobasal) segment longest, slightly longer than penultimate seg- ment. Labial palpus small (Fig. 3), 3 segmented; first segment shortest, about half as long as second segment; second segment about as long as ultimate segment, with a single long yellow seta apicodorsally; ultimate segment broadest, compressed laterally, bearing a single long yellow seta on apicoventral angle. Pronotum slightly more than twice as wide as long; widely margined lat- erally, finely margined anteriorly, and very finely posteriorly; punctures slightly coarser than those on head and separated by one-half to one times their width except an anterolateral and a mediolateral series of very coarse punctures; anterolateral and posterolateral angles rounded. Prosternum non- carinate (Fig. 2). Elytron convex, distinctly margined laterally, widest slightly behind mid- length. Surface with punctures finer and sparser than those of pronotum except three incomplete serial rows of coarser punctures discally, one sim- ilar but more distinct lateral row, and rudiments of another row along elytral margin. Sutural stria present from apex to about basal fourth. Epipleura strongly vertical. Scutellum an equal-sided triangle. Mesosternum finely alutaceous; with a fine, very low longitudinal carina on posterior half and extending between mid-coxae (Fig. 4). Metasternum finely alutaceous and punctate except a small glabrous swollen area on mid- line in front of inner angles of hind coxae and across hind margin adjacent to hind coxae. Front, middle, and hind femora densely covered with short hydrofuge pubescence on basal three-fourths, apices glabrous. Metatrochanters not elongated. Metatibiae not arcuate. Tarsal formula 5-5-5. Foreleg with seg- ments | to 4 about equal in length; last segment about as long as segments 1 to 4 combined. Midleg and hindleg both with basal segment short, ter- minating obliquely apically; second segment longer than third segment; third and fourth segments shortest and subequal; fifth segment slightly longer than third and fourth segments combined. Abdominal sterna finely aluta- ceous; covered with moderately dense, short, yellowish hydrofuge pubes- cence; last sternum with very shallow apicomedial notch (Fig. 5). Genitalia as illustrated (Figs. 6, 7). Allotype.—Similar to male. Variations.—Males vary in length from 2.8 mm to 3.3 mm; females vary VOLUME 92, NUMBER 4 Fig. 2. Dieroxenus cremnobates, n. sp., head and appendages, ventral view; prosternum, 80x. Fig. 3. Dieroxenus cremnobates, n. sp., mentum and labial palpi, 210x. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 4. Dieroxenus cremnobates, n. sp., mesosternum, oblique view, head to left, 190x. Fig. 5. Dieroxenus cremnobates, n. sp., last abdominal sternum, 500x. VOLUME 92, NUMBER 4 759 0.5mm Figs. 6-7. Dieroxenus cremnobates, n. sp., male genitalia: 6, dorsal view: 7, ventral view. from 2.7 mm to 3.5 mm. In teneral and mature specimens the head is pi- ceous, however, in teneral specimens the pronotum and elytra are uniformly reddish brown instead of broadly piceous on discal areas. Type-data.—Holotype: ECUADOR, Napo, Baeza (72 km E), 16 May 1975, Spangler, Langley, and Cohen; USNM Type No. 73590, deposited in the National Museum of Natural History, Smithsonian Institution. Allotype: same data as holotype. Paratypes: same data as holotype, 44 6d, 42 29; ECUADOR, Tunguruhua: Banos (35 km E), 29 May 1975, Langley, Cohen, and Monnig, 8 64,6 2 2; Banos (18 km E), 25 Jan. 1976, Spangler, et al., 15 66, 18 22; Banos (20 km E), 28 Jan. 1976, Spangler et al., 7 dd, 6 2 2; Banos (39 km E), 28 Jan. 1976, Spangler et al., 2 °°. Paratypes will be deposited in the following entomological collections: American Museum of Natural History, New York; British Museum (Natural History), London; California Academy of Sciences, San Francisco; Cana- dian National Collection, Ottawa, Canada; Institut fur Pflanzanschutzfor- schung Zweigstelle, Eberswalde, East Germany; Museo Argentino de Cien- cias Naturales, ‘‘Bernardino Rivadavia,’’ Buenos Aires, Argentina; Museum of Comparative Zoology, Cambridge; Museo Nacional de Historia 760 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 8.. Dieroxenus cremnobates, n. sp., biotope. Natural, Santiago, Chile; Museum fiir Naturkunde, East Berlin, Deutsche Demokratische Republik; Museum National d’Histoire Naturelle, Paris, France; Institut Royal des Sciences Naturelles de Belgique, Bruxelles, Bel- gique; and Zoologische Sammlung Bayerischen Staates, Munchen, West Germany. Etymology.—cremnobates from kremnobates, G.—frequenter of steep places in reference to the nearly vertical aquatic habitats on which these beetles were found. Habitat.—The type-specimens were found on seeps, i.e., on the wet sur- faces of rocks and in crevices of these rocks which were exposed by road cuts. The habitat at 18 km east of Banos is illustrated in the photograph (Fig. 8). Specimens of this new genus were found intermixed with members of the hydrophilid genera Oocyclus and Anacaena on the seeps. Acknowledgments The first specimens of this new genus were collected by P. J. Spangler, Andrea Langley and Jeffrey Cohen in 1975 during fieldwork in South Amer- ica supported by a Smithsonian Institution Fluid Research Grant used to initiate a combined Ecuador-Peace Corps-Smithsonian Institution Aquatic VOLUME 92, NUMBER 4 761 Insect Survey of Ecuador. Additional specimens were collected in 1976 through the auspices of a grant from the National Geographic Society. I am very grateful to these sponsors for their support. I also extend my thanks to Mrs. Elaine Hodges, Smithsonian Institution staff artist, for preparing the illustrations; to Mrs. Suzanne Braden, Smith- sonian Institution scanning electron microscopist, for taking the micro- graphs for this article; and to Andrea Langley-Armstrong and Jeffrey Cohen, former Peace Corps Volunteers and Paul Monnig for their collecting efforts which provided some of the paratypes of this new genus as well as many other aquatic insects from Ecuador. Reference Cited d’Orchymont, A. 1942. Contribution a |’etude de la Tribu Hydrobiini Bedel Sous-Tribu Hy- drobiae (Palpicornia-Hydrophilidae).—Musee Royal d’Histoire naturelle de Belgique, 2nd series, Fascicle 24:1-68. Smithsonian Institution, Washington, D.C. 20560. PROC. BIOL. SOC. WASH. 92(4), 1979, pp. 762-768 A REDESCRIPTION OF SCOLOPLOS SIMPLEX (HUTCHINGS, 1974) (POLYCHAETA: ORBINIIDAE) FROM AUSTRALIA Betsy Brown Abstract.—The orbiniid polychaete Scoloplos simplex (Hutchings, 1974) from Australia is redescribed and compared to Scoloplos fragilis (Vermill, 1873) from the east coast of North America. The genera Scoloplos Blain- ville, Haploscoloplos Monro and Leitoscoloplos Day are discussed. In connection with a study on the distribution of Scoloplos fragilis (Ver- rill, 1873) in the intertidal zone of a geologically-active tidal flat in Delaware Bay (Brown, 1979), I reviewed the literature on the biology, reproduction and geographic distribution of the species. In her study on the polychaetes of the New England region, Pettibone (1963:292) reported that S. fragilis was known from the Gulf of St. Lawrence and Newfoundland to Florida and the Gulf of Mexico, in low water to 102 meters. Anderson (1960: 105) identified some specimens from the sand flats of Botany Bay, New South Wales as Haploscoloplos fragilis (see discussion of Scoloplos, Haploscol- oplos, and Leitoscoloplos below). In the following year, Anderson (1961:257) reported on the embryonic and postembryonic development of this Australian species from Botany Bay. These records of H. fragilis from New South Wales were not mentioned by Pettibone (1963) since they ap- peared while her paper was in press (Pettibone, in correspondence). Ref- erence to the development of H. fragilis by Anderson was cited by Schroe- der and Hermans (1975:130) in their recent review of polychaete reproduction. Using the latter publication, Holland and Polgar (1976:346) summarized the reproductive mechanism of H. fragilis (Holland, personal communication). To my knowledge, the development of S. fragilis from the east coast of North America has yet to be described. It should be noted here that the two references of Anderson (1960, 1961) were not mentioned by Hutchings (1974:183) when she described a new species of Haploscolo- plos from New South Wales or by Day (1977:217) in his review of the Australian and New Zealand Orbiniidae. Thus it was of interest to examine some of Anderson’s specimens from Botany Bay and compare them with my specimens of S. fragilis from Del- aware Bay. Anderson (1960:105) mentioned that his specimens differed from the description of H. fragilis by Hartman ‘‘only in the absence of an inter- ramal cirrus on the anterior abdominal parapodia, a character very variable VOLUME 92, NUMBER 4 763 within the family.’’ In correspondence with Dr. Pettibone, it was learned that Dr. Anderson’s collection of H. fragilis from Botany Bay was not available in the United States National Museum of Natural History or in the Australian Museum (information from Dr. P. Hutchings). Dr. Anderson rec- tified the situation by very kindly collecting some additional specimens from the site in Botany Bay where he had made his previous collections and carried out his developmental studies. Seventeen specimens (USNM 54834) were sent to Dr. Pettibone and then forwarded to me on loan for study. They proved to belong not to S. fragilis (Verrill) but to Haploscoloplos simplex Hutchings (1974:183), described from Wallis Lake, New South Wales and referred to Scoloplos (Scoloplos) simplex (Hutchings) by Day (1977:228), who added additional records from Australia: Queensland, New South Wales, Victoria and Western Australia. A paratype of H. simplex (USNM 49487) was also available for study. The specimens of S. simplex from New South Wales are described below and compared with S. fragilis from the east coast of North America, followed by a discussion of Scoloplos Blainville, Haploscoloplos Monro, and Leitoscoloplos Day. FAMILY ORBINIIDAE (=ARICIIDAE) Scoloplos Blainville Scoloplos simplex (Hutchings) Haploscoloplos fragilis.—Anderson, 1960:105—106; 1961:257—272, figs. 1, 2.—Schroeder and Hermans, 1975:130 (data from Anderson, 1961). Not Verrill, 1873. Haploscoloplos simplex.—Hutchings, 1974:183-—184, fig. 2A—D. Scoloplos (Scoloplos) simplex.—Day, 1977:228—229. Material examined.—New South Wales, Australia: Botany Bay, January 1977, D. T. Anderson, coll.—17 specimens (USNM 54834). Wallis Lake, sta. 401-450, sandy, December 1970, Univ. of N.S.W., coll.—paratype of Haploscoloplos simplex (USNM 49487). Description.—The prostomium is conical, longer than wide when com- plete (Fig. 1a). Deep slits, one located anteriolaterally on each side of the achaetous, single-ringed peristomium, form the nuchal organs. No eyespots are visible. The proboscis is eversible and foliose (Hutchings, 1974, Fig. 2A). The width, measured at the junction between the thorax and abdomen, ranges from 1.4 to 2.6 mm. Branchiae appear first as small papillae on setigers 9-12 (Table 1). This character does not depend on the size of the organism. In most cases, branchiae become fully developed on the first abdominal setiger. Occasionally, the transition between the thorax and ab- domen occurs over three setigers with the intermediate setiger having bran- chiae intermediate between the papillate and well-developed forms. The transition between thorax and abdomen occurs more posteriorly in larger 764 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 1. Scoloplos simplex: a, Anterior end, lateral view; b, Parapodia of 4th setiger, an- terior view; c, Crenulate capillary setae; d, Thoracic, neuropodial hook. VOLUME 92, NUMBER 4 765 Table 1.—Morphological and sexual variability among specimens of Scoloplos simplex (Hutchings). Setiger** Setiger # of Specimen Width* branchial transition between number (mm) papillae begin thorax and abdemen Sex 1 1.4 9 14-15 Male Z 1.4 (Mais, 14-15 Male 3 1.6 10, 11 15-16 Y - 1.6 11 14-15 Male 5 i 9 14-16 Male 6 eZ 10 14-15 Male ih 1.8 1] 14-16 Male 8 1.8 est 14-15 ? 9 2.0 12 14-16 Female 10 2.0 12 14-15 Female 11 2:2) 10 14-15 Female 12 De3 9, 10 15-16 Female 13 (paratype) P2283 9.10 16-17 Female 14 2.4 11 16-17 Male LS ieee 12 16-17 Female 16 2.6 oF 10 16-17 7 17 2.6 10 16-17 Female 18 2.6 10 16-17 Male * Measured at the junction between the thorax and abdomen. ** The branchial papillae often begin on different setigers on opposite sides of the same specimen. Numbers indicate the setiger on which the papillae begin on the opposite sides of the specimen. The anterior-most papilla is listed first. specimens (Table 1) and appearance of the first abdominal setiger ranges from setiger 15 to setiger 17 (up to setiger 18, according to Hutchings). Thoracic notopodia and neuropodia have simple, papillate postsetal lobes (Fig. 1b), which become increasingly cirriform towards the posterior end of the thorax. Thoracic notosetae are all crenulate capillaries (Fig. lc). Two types of thoracic neurosetae are found: 1) crenulate capillaries (Figs. 1b, c) and 2) short hooks, 6—8 per neuropodia, with slightly bent tips and present on all but the last one or two setigers of the thorax (Fig. 1d). Abdominal setae are all crenulate capillaries. In the abdomen, the cirniform, postsetal notopodial lobes are equally as long as the notosetae and branchiae. The branchiae and abdominal notopodia emerge dorsally. The abdominal neu- ropodia emerge dorsolaterally and are bilobed, with inner lobes longer than outer lobes. The outer lobe is incised just below its tip (Fig. la), below which the ventrolateral margins may have a whitened, glandular appear- ance. Anal cirri are absent. In most cases, sex could be determined, with females appearing to be larger than males (Table 1). Preserved specimens are white or buff colored. 766 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Distribution.—Along coast of Queensland, New South Wales, Victoria and Western Australia. Remarks.—The above description differs slightly from that of Hutchings (1974). Hutchings noted, “Faded eyespots are arranged in two oval patches at the base of the prostomium.’’ The nuchal organs, described above, are situated in oval patches at the base of the prostomium and are probably equivalent to Hutchings’ eyespots. Two types of thoracic neurosetae were noted by Hutchings: **(1) long distally pointed spinous setae, which when viewed side on have a toothed appearance, and (2) 10-15 simple acicular type setae with a slightly bent tip. The bases of these setae appear to be split, and they are slightly more chitinized than the spinous setae.’ The neuropodial, crenulate capillaries observed in this study are synonymous with Hutchings’ type 1 “‘spinous setae.’’ The hooks seen in this study, probably equivalent to Hutchings’ type 2 “‘acicular type setae,’’ were fewer in number (6—8 per bundle) than those observed by Hutchings (10-15 per bundle), indicating considerable variation in this character. The bases of the hooks observed in this study did not appear to be split. Scoloplos simplex (Hutchings) from Australia differs from Scoloplos fra- gilis (Verrill) from the east coast of North America in the following ways: S. simplex S. fragilis Thoracic neurosetae Short hooks and Long capillaries long capillaries only Interramal cirri on anterior Absent Present abdominal segments Anal cirri Absent Present Subpodal papillae at Absent Present junction of thorax and abdomen Discussion of Scoloplos Blainville, Haploscoloplos Monro and Leito- scoloplos Day: Scoloplos Blainville (1828:493), with type-species Lumbri- cus armiger O. F. Miller by monotypy, is characterized by a pointed pro- stomium and a one-ringed, achaetous buccal segment or peristomium, the thoracic neuropodia have unfimbriated, postsetal lobes with 0-3 papillae, sometimes with subpodal or ventral papillae on some segments; the thoracic neurosetae include crenulate capillaries and sometimes blunt hooks. Monro (1933:261) established the genus Haploscoloplos, with type-species Scolo- plos cylindrifer Ehlers (1905) by original designation, with the diagnosis: ‘“As Scoloplos but with crenate capillary bristles only.’’ Hartman (1944:340, 1957:269) retained the distinction between the two genera, while Pettibone (1957:160) synonymized them stating, ‘‘In some cases this character is dif- ficult to observe; the crotchets may be easily missed, occurring in some thoracic neuropodia and not in others. Until it can be established how much the abrasive action of certain substrate has to do with the formation of VOLUME 92, NUMBER 4 767 certain types of crotchets from capillaries, the character does not seem a good one.”’ Curtis (1969:3280) presented evidence from benthic samples that sug- gested hooks appeared in Scoloplos armiger as it matured and were lacking in juveniles. I gathered similar data on S. armiger from George’s Bank off Massachusetts. In agreement with Pettibone, Curtis was of the opinion that the absence of hooks is not a sufficient criterion for generic distinction. In a review of the subfamily Orbiniinae, Day (1973:86) stated, *‘To me there seems no possibility that the normal hooks with rounded ends and guards could have been formed from broken or abraded crenulate capillaries. While I recognize that juvenile specimens of some species of Scoloplos may be wrongly assigned to Haploscoloplos, 1 agree with Hartman and Monro in recognizing it as a valid genus.’’ Both genera have continued to be used to describe species without hooks in the thoracic neuropodia. In connection with a review of the Australian and New Zealand Orbini- idae, Day (1977:218) examined a specimen of Scoloplos cylindrifer Ehlers, the type-species of Haploscoloplos as selected by Monro, and found on examination of the thoracic neurosetae ‘‘an anterior row of short, slender hooks that is easily overlooked. Thus the type species of Haploscoloplos is a synonym of Scoloplos.’’ For the species that did fit the definition of Haploscoloplos, Day (1977:218) erected a new genus Leitoscoloplos. The small thoracic neuropodial hooks in Scoloplos cylindrifer had previously been overlooked by Ehlers (1905:45), Monro (1939:124), and many others. The short rows of thoracic neuropodial hooks in the orbiniids from New South Wales, identified as Haploscoloplos fragilis by Anderson (1960, 1961), were also overlooked. In order to designate a new genus, the separating character should be readily detectable and clear cut in its segregation of the group possessing it (Blackwelder, 1967:203; Mayr, 1969:92). Use of any single character is less likely to give satisfactory results in natural classification than if the character correlates with other characters descriptive of the group (Black- welder, 1967:203). The use of the presence or absence of hooks to separate Leitoscoloplos or Haploscoloplos from Scoloplos is not a readily detectable character. Accordingly, the synonymy of Leitoscoloplos and Haploscolo- plos with Scoloplos has been retained here. Acknowledgments I am grateful to Dr. Anderson for collecting and contributing specimens from Australia. Robert Palmer and Robert Prezant provided useful criticism of the manuscript during preparation. I am especially grateful to Dr. Marian H. Pettibone of the Smithsonian Institution for the loan of specimens and for critically reviewing the manuscript. This is contribution 147, University of Delaware, College of Marine Studies. 768 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Literature Cited Anderson, D. T. 1960. Ariciid polychaetes in Australia.—Proc. Linn. Soc. N.S.W. 85:105- 106. . 1961. The development of the polychaete Haploscoloplos fragilis.—Quart. J. Microsc. Sci. 102(2):257—272. Blackwelder, Richard E. 1967. Taxonomy.—John Wiley & Sons, Inc., N.Y., 608 pp. Blainville, Henri de. 1828. Jn Dictionnaire des Sciences Naturelles, 57:368—501. Brown, Betsy. 1979. The distribution and biology of Scoloplos fragilis (Orbiniidae: Poly- chaeta) on a tidal flat, Cape Henlopen, Lewes, Delaware.—M.S. Thesis, University of Delaware. 115 pp. Curtis, Mark A. 1969. Synonymy of the polychaete Scoloplos acutus with S. armiger.—J. Fish. Res. Bd. Canada 26(12):3279-3282. Day, J. H. 1973. New Polychaeta from Beaufort, with a key to all species recorded from North Carolina.—NOAA Tech. Rept. NMFS Circ-375, 140 pp. . 1977. A review of the Australian and New Zealand Orbiniidae (Annelida: Polychae- ta).—Pp. 217-246 in Donald J. Reish and Kristian Fauchald, eds. Essays on Polychae- tous Annelids in Memory of Dr. Olga Hartman. Special Publication, Allan Hancock Foundation. Ehlers, E. 1905 (1904). Neuseelandische Anneliden.—Abh. K. Gesell. Wiss. Gottingen, Math.-Phys. KI. N.F. 3(1):1-80. Hartman, Olga. 1944. New England Annelida. Pt. 2, including the unpublished plates by Vernill with reconstructed captions.—Bull. American Mus. Nat. Hist. 82(7):327-344, pls. 13-35, 45-60. . 1957. Orbiniidae, Apistobranchidae, Paraonidae, and Longosomidae.—Allan Hancock Pacific Exped. 15(3):1-393. Holland, A. F., and T. T. Polgar. 1976. Seasonal changes in the structure of an intertidal community.—Mar. Biol. 37:341-348. Hutchings, Patricia. 1974. Polychaeta of Wallis Lake, New South Wales.—Proc. Linn. Soc. of New S. Wales 98(4): 175-195. Mayr, Ernst. 1969. Principles of systematic zoology.—McGraw-Hill Book Company, N.Y., 428 pp. Monro, C.C. A. 1933. Onacollection of Polychaeta from Dry Tortugas, Florida.—Ann. Mag. Nat. Hist., ser. 10, 12:244—269. 1939. Polychaeta.—Reports B.A.N.Z. Antarctic Research Expedition, 1929-1931, ser. B, 4(4):87-156. Pettibone, Marian. 1957. North American genera of the family Orbiniidae (Annelida: Poly- chaeta), with descriptions of new species.—J. Wash. Acad. Sci. 47:159-167. . 1963. Marine polychaete worms of the New England region. I. Families Aphroditidae through Trochochaetidae.—U.S. Natl. Mus. Bull. 227(1): 1-356. Schroeder, Paul C., and Colin O. Hermans. 1975. Ch. 1 Annelida: Polychaeta.—Pp. 1-213 in Arthur C. Giese and John S. Pearse, eds., Reproduction of marine invertebrates. vol. 3. Verrill, Addison Emory. 1873. Report upon the invertebrate animals of Vineyard Sound and the adjacent waters, with an account of the physical characters of the region.—Rep. U.S. Fish. Comm. for 1871-72:295-852, 39 pls. College of Marine Studies, University of Delaware, Lewes, Delaware 19958. PROC. BIOL. SOC. WASH. 92(4), 1979, pp. 769-772 A NEW GENUS AND SPECIES OF BOPYRID ISOPOD INFESTING THE CRANGONID SHRIMP PONTOPHILUS ABYSSI SMITH IN DEEP WATER OF THE NORTHWESTERN ATLANTIC OCEAN John C. Markham Abstract.—Pontobopyrus abyssorum, a new genus and species of pseu- dionine bopyrid isopod branchially infesting the crangonid shrimp Ponto- philus abyssi Smith from a depth of 3850 m off the coast of Virginia, is described. This depth is evidently the greatest known for the occurrence of any species of bopyrid. The new species is markedly different from all others known to infest caridean shrimps. A single female bopyrid isopod from the branchial chamber of the cran- gonid shrimp Pontophilus abyssi S. 1. Smith has become available. It clearly represents a hitherto undescribed genus and species of the bopyrid subfam- ily Pseudioninae. Regrettably no male was obtained, but because it was unlikely that one would be found, it seemed appropriate to prepare the description without it. Pontobopyrus, new genus Diagnosis. —Female: Body axis describing nearly circular arc; all body segments set apart. Head deeply set into pereon and completely bordered anteriorly by large frontal lamina; maxilliped with articulating palp; pos- teroventral border of head bearing 2 pairs of lateral projections. Oostegites completely enclosing brood pouch. Six pleomeres deeply separated later- ally, first 5 produced into elongate lateral plates and bearing small uniramous pleopods; final pleomere extended into long uniramous uropods. Male un- known. Etymology and gender.—Prefix “‘ponto-’’ from Greek word meaning ‘‘open sea’’ in reference to occurrence in deep ocean and in reflection of host’s generic name Pontophilus, combined with Bopyrus, name of type- genus of family Bopyridae. Gender masculine. Type-species.—Pontobopyrus abyssorum, new species. Pontobopyrus abyssorum, new species Fig. 1 Material examined.—Infesting Pontophilus abyssi S. I. Smith. Cruise 770 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 1. Pontobopyrus abyssorum, holotype female: a, Dorsal view; b, Ventral view; c, Right antenna 1; d, Right antenna 2; e, Right maxilliped; f, Palp of right maxilliped; g, Spur of right maxilliped; h, Right posteroventral border of head; i, Left oostegite 1; external; j, Same, internal; k, Right pereopod 1; 1, Right pereopod 7. Scale line: 4.0 mm for a, b, i, j; 2.0 mm for e, h; 1.8 mm for k, 1; 0.7 mm for c, d, f, g. EPA-78-01 of R/V Advance II, Station 9: Northwestern Atlantic Ocean, 38°04'14"N to 38°09'15”N, 70°26'22"”W to 70°22'10”W, charted depth 3,850 m, 24 June 1978, 3-hour tow with 45-foot semi-balloon otter trawl, J. A. Musick and K. Sulak, colls., E. L. Wenner, det. of host. 1 2, holotype, USNM 172350. Description of holotype female (Fig. 1).—Length (exclusive of uropods) 9.76 mm, maximal width 5.40 mm, head length (exclusive of frontal lamina) 1.01 mm, pleon length (exclusive of uropods) 3.23 mm. Body axis evenly and continuously distorted along nearly circular arc through about 77°. All body regions and segments distinct (Fig. la, b). Head subelliptical, much broader than long, deeply set into pereon. Large fleshy frontal lamina reflexed over whole anterior margin. No eyes. First antenna (Fig. Ic) of 3 articles rapidly decreasing in size distally; second antenna (Fig. Id) of 4 articles also much smaller distally; distal articles of both possibly setose, but not discernibly so. Maxilliped (Fig. le) quite long, with irregular border; prominent acutely pointed setose palp arising from VOLUME 92, NUMBER 4 771 anterior margin about 3 of distance from anteromedial corner, distinctly articulating with maxilliped; anteromedial corner of posterior maxillipedal segment produced into well-developed setose spur (Fig. 1g). Posteroventral border of head (Fig. 1h) with 2 rather widely spaced clavate lateral projec- tions on each side, nearly entire margin in middle. Pereomeres deeply separated, especially on long side. Coxal plates on both sides of pereomeres 1-4; tergal projections on both sides of all per- eomeres, though indistinctly set off posteriorly. Oostegites arching high and completely enclosing brood pouch; first oostegite (Fig. li, j) rather small, relatively narrow, produced into broad blunt posterolateral point, its internal ridge unadorned; other oostegites large, each about twice as wide as long. Pereopods (Fig. 1k, 1) all of about same size, but basal articles of anterior ones much larger; some meri and carpi fused. Pleon extended, of 6 pleomeres all deeply divided laterally. Pleomeres 1- 5 developed into sharply tapered lanceolate lateral plates on both sides, those on longer side considerably larger. Five pairs of short posteriorly directed uniramous lanceolate pleopods along sides but not at all concealing middle of pleon. Final pleomere produced into long slender tapering uni- ramous uropods, their length exceeding half of rest of length of pleon. Male unknown. Etymology.—Specific name abyssorum, ‘‘of the deeps,’’ genitive plural of host’s specific name and indicator of depth of collection of species. Discussion.—Although clearly assignable to the subfamily Pseudioninae (Codreanu, 1967), the female of Pontobopyrus abyssorum is quite different from that of any genus heretofore described in that subfamily. The enclosure of the head is highly unusual, more reminiscent of the condition in the Orbioninae. The shapes of the tergal projections, oostegites and lateral plates and the body curvature are also distinctive. Finally, the combination of an articulating maxilliped palp, uniramous pleopods and elongate unira- mous uropods immediately distinguishes it from other pseudionine genera. The subfamily Pseudioninae contains many species, mostly parasites of anomurans, but a few species of Pseudione are known to infest carideans, especially crangonids (including 2 species of Pontophilus), pandalids and nephropids. (See Bourdon, 1968, 1971, for partial listing.) A thorough re- vision of the ill-defined genus Pseudione will probably result in the reas- signment of some of the caridean-infesting species to new genera, but no other species of pseudionine recorded as the parasite of a caridean is so obviously not a member of Pseudione as Pontobopyrus abyssorum. Other branchial bopyrid parasites of carideans include the whole of the large subfamily Bopyrinae (Codreanu, 1967) and most of the small subfamily Ar- gelinae (Markham, 1977). Evidently the deepest a bopyrid has been recorded is a depth of 800 to 900 fathoms (=1,460-1,650 m) for the appropriately named species Bathy- 772 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON gyge grandis Hansen, infesting Glyphocrangon sculptus (Smith) off South Africa (Stebbing, 1908). That is less than half the record for Pontobopyrus abyssorum, at 3,850 m. Acknowledgments Sincerest thanks are due to Elizabeth Lewis Wenner of the South Carolina Wildlife and Marine Resources Department for making the type-specimen available for study and for furnishing collection data for it. This report was prepared under National Science Foundation Grant Number 76-20102, ad- ministered by the Bermuda Biological Station for Research, Inc., of which this is Scientific Contribution Number 815. Literature Cited Bourdon, R. 1968. Les Bopyridae des mers Européenes.—Mem. Mus. Natl. Hist. Nat. (Paris), N. S. (A) 50 (2):77-424. . 1971. Epicarides nouveaux pour la c6te occidental d’ Afrique équatoriale.—Bull. Inst. Fond. Afrique Noire (A) 33:371-391. Codreanu, R. 1967. Clasificarea evolutiva a bopirienilor, isopode parazite ale crustaceelor decapode si importanta lor biologica generala.—Stud. Cerc. Biol. (Ser. Zool.) (Bucarest) 19:203-211. Markham, J. C. 1977. Description of a new western Atlantic species of Argeia Dana with a proposed new subfamily for this and related genera (Crustacea Isopoda, Bopyridae).— Zool. Meded. 52:107—123. Stebbing, T. R. R. 1908. South African Crustacea (Part IV).—Ann. South African Mus. 6: 1-96. Bermuda Biological Station for Research, Inc., St. George’s West, 1-15, Bermuda. Present address: Arch Cape Marine Laboratory, Arch Cape, Oregon 97102, U.S.A. PROC. BIOL. SOC. WASH. 92(4), 1979, pp. 773-782 RECTIFICATIONS IN THE NOMENCLATURE OF SOME INDO-PACIFIC LITTORINIDAE Winston F. Ponder and Joseph Rosewater Abstract.—Three species previously placed in the subgenus Littoraria Griffith and Pidgeon, 1834, are reassigned to Austrolittorina Rosewater, 1970. Several other nomenclatorial inconsistencies involving species of Indo-Pacific Littorinidae are clarified. New information which has become available since the publication by Rosewater (1970) of a classification of Indo-Pacific Littorinidae, subfamily Littorininae, has resulted in changes in the subgeneric assignments of cer- tain species. Assignments to subgenera had then to be made subjectively in some cases because of the lack of anatomical material, and decisions as to placement were based mainly on shell characteristics. As animals have now become available and have been examined with special reference to the radulae and male reproductive anatomy, certain changes have been deemed necessary. The opportunity is also taken to rectify the taxonomy of two small Aus- tralian littorinids not included in Rosewater’s (1970) revision: Laevilitorina jJohnstoni (Cotton, 1945), and L. mariae (T. Woods, 1876). In addition several other changes are brought to light, all of which are summarized in Table 1. Littorina (Austrolittorina) acutispira E. A. Smith, 1892. Pii-l.-hes..1—4:- PE 2, fis. Littorina acutispira E. A. Smith, 1892:487, pl. 40, fig. 3.—Rosewater, 1970:451, pl. 349, fig. 3; pl. 351, fig. 1. Types: 14 syntypes (B.M.N.H.’, 1891.11.6.216—225). Rock pools, Green Point, Watson Bay, Port Jackson, New South Wales. Littorina infans E. A. Smith, 1892:488, pl. 40, fig. 4; Rosewater, 1970:452, pl. 351, fig. 2. Types:—14 syntypes (B.M.N.H., 1891.11.6.226—235). Green Point, Watson Bay, Port Jackson, New South Wales. Remarks.—Rosewater (1970) tentatively placed L. acutispira in the sub- genus Littoraria Griffith and Pidgeon, 1834, and L. infans was considered a doubtful member of the Littorinidae. Large samples collected in the vi- 1 British Museum (Natural History), London. 774 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Table 1. Former assignment Littorina (Littoraria?) acutispira E. A. Smith, 1892 [Rosewater, 1970] Littorina (Littoraria?) infans E. A. Smith, 1892 [Rosewater, 1970] Littorina (Littoraria) praetermissa May, 1909 [Rosewater, 1970] Littorina paludinella Reeve, 1857 [Macpherson and Gabriel, 1962] Littorina (Littoraria?) sundaica Altena, 1945 [Rosewater, 1970] Pellax johnstoni Cotton, 1945 Laevilitorina burni Ponder, 1976 Rissolittorina mariae (T. Woods, 1876) [Ponder, 1966] Littorina hisseyiana T. Woods, 1876 [=Trochacea, Rosewater, 1970] Littorina (Littorinopsis) incisa Yokoyama, 1927 [Rosewater, 1970] Littorina (Littoraria) lucida Corrected assignment Littorina (Austrolittorina) acutispira E. A. Smith, 1892 Littorina (Austrolittorina) praetermissa May, 1909 Littorina (Austrolittorina) sundaica Altena, 1945 Laevilitorina johnstoni (Cotton, 1945) Laevilitorina mariae (T. Woods, 1876) Hisseyagibbula hisseyiana (1. Woods, 1876) Pyramidellidae Assiminea japonica Martens, 1877 Yokoyama, 1927 [Rosewater, 1970] cinity of Sydney (the type-locality of both species) has shown that these two names are based on forms of one species. The types of L. acutispira are tall-spired, clean shells whereas those of L. infans are smaller, stunted, eroded specimens. Both species names were introduced simultaneously, and we, as first revisers, select L. acutispira Smith as the valid name to be used. The lectotype of L. acutispira is a more mature and better preserved indi- vidual so that there is less chance of confusion in its future identification. It is also the name recognised by Rosewater (1970) for this species, although he considered its subgeneric classification to be doubtful. The radula (Pl. 2, fig. 5) and penis (Pl. 1, fig. 4) are like those of species of the subgenus Austrolittorina Rosewater, 1970. The shell coloration is similar to that of L. praetermissa, a species shown herein to also belong to Austrolittorina. Three specimens are figured to illustrate the range of variation of this species (Ei ines. l—3)) This species is found in southern Queensland, New South Wales and westernmost Victoria. It is very abundant over most of its range, living together with L. unifasciata Gray, 1826. VOLUME 92, NUMBER 4 775 Littorina (Austrolittorina) praetermissa May, 1909 Pl. 33.812... 3 Litorina praetermissa May, 1909:57, pl. 6, fig. 3. Type: Tasmanian Museum, Hobart, Tasmania. Remarks.—Rosewater (1970) eliminated from consideration as a littorinid taxon the name Littorina paludinella Reeve, 1857, used by a few early Tasmanian and Victorian workers (T. Woods, 1877:36; Pritchard and Gatliff, 1902:92) because Hedley (1913:283) had already concluded that Reeve’s species is a Tasmanian hydrobiid and an earlier name for Ampullaria tas- manica T. Woods, 1876, and that the littorinid, sometimes called L. palu- dinella is the juvenile of L. praetermissa. More recently Macpherson and Gabriel (1962:89; Macpherson, 1966) have used the name Littorina paludi- nella for a small, dark littorinid, but most subsequent authorities have fol- lowed Hedley’s conclusion. Macpherson and Gabriel (ibid.) stated that ‘*Hedley confused this species with Ampullaria tasmanica Tenison Woods, 1876, a Tasmanian fluviatile species without the flared lip.’’ Reference to Reeve’s specimens in the British Museum (N.H.) (also examined by Hedley) and to his original figure, show that Hedley’s conclusion is the only reason- able one. Macpherson and Gabriel (ibid.) noted that Littorina paludinella also extends into New South Wales, but as L. praetermissa does not occur in that state they were probably confusing it with L. acutispira. The juveniles of L. praetermissa are more uniformly black and more globular than L. acutispira and in some localities mostly occur low on the shore suggesting that at least part of the population undergoes an up-shore migration from about the lower mid-littoral zone to the upper littoral where the adults are normally found. A number of samples were examined and the small *‘pal- udinella’’ form of L. praetermissa were all found to be immature. This species lives sympatrically with L. (Austrolittorina) unifasciata in Tasman- ia, Victoria and South Australia, where they coexist in the upper littoral zone. L. praetermissa and L. acutispira do not appear to overlap in distri- bution. Rosewater (1970) placed L. praetermissa in the subgenus Littoraria, main- ly on the basis of penial morphology which was deduced from poorly pre- served material. He remarked that the radula is similar to that of L. (Aus- trolittorina) unifasciata and that there is an umbilical crescent present, although not as deeply impressed as that of L. (Austrolittorina) unifasciata. Both of these latter characters indicate placement in the subgenus Austro- littorina Rosewater and re-examination of the penis of L. praetermissa sup- ports this view as it is nearly identical to that of L. (Austrolittorina) unifas- ciata (Pl. 3, fig. 3). PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 776 02 mm VOLUME 92, NUMBER 4 777 Littorina (Austrolittorina) sundaica Altena, 1945 Pl. 2, figs. 1-4; Pl. 3, figs. 1, 2 Littorina (Melaraphe) sundaica Altena, 1945:151, fig. 2. Type: Holotype (Rijksmuseum van Natuurlijke Historie, Leiden, no. 52038. Tjilaoet Eureun, South Coast of Java). Remarks.—Rosewater (1970) placed Littorina sundaica Altena, 1945, in the subgenus Littoraria which he characterized as having a penis with a basal flap but no apparent accessory glands. The members of the subgenus were said not to have white apertural bands, a notable characteristic of L. sun- daica which should have eliminated it. It was placed in Littoraria on the basis of shell characteristics but questionably, nevertheless, pending ana- tomical evidence. Preserved specimens only recently received from Arie Budiman, Museum Zoologicum Bogoriense, which he collected at Pangandaran, Java, have permitted examination of the radula and penis of this species (Pl. 3, figs. i 32) Several characteristics quite clearly relate L. sundaica to the subgenus Austrolittorina, including a penis with a basal enlargement containing a penial gland and accessory flagellum (Rosewater, 1970:467) and a radula with a rather narrow central tooth. The flattened columella with crescent- shaped area on adjacent shell that is characteristic of the type-species of Austrolittorina, L. unifasciata, is missing or reduced in some other mem- bers of the subgenus including L. sundaica. The several specimens from Java and another specimen from Bali also change somewhat the picture of the species gained from the few specimens available previously. Of 15 specimens measured, the lengths varied from 4.5 mm to 7.2 mm, widths from 2.4 to 3.6 mm, and the average obesity of the specimens was .52 compared with .55 in the 4 specimens available pre- viously (Rosewater, 1970). New localities for the species are: Pangandaran (on the mid to western middle of the south coast), Java (A. Budiman, 1976; USNMz?) and Sanur, Bali (F. G. Thompson, 1971, USNM). Sexual dimorphism does not seem strikingly apparent in the shells of the 2 National Museum of Natural History, Washington. << Plate 1. Figs. 1-4. Littorina (Austrolittorina) acutispira: 1-3, Shells to show range of variation; from Bottle and Glass Rocks, Vaucluse, Sydney, New South Wales, Australia (1) 6.0 x 3.2 mm (2) 4.3 < 2.6 mm (3) 3.8 X 2.6 mm. (Australian Museum, C.114545); 4a, b, Penis from adult specimen collected at Edwards Beach, Balmoral, Sydney, New South Wales, Aus- tralia, Nov. 1978 (Australian Museum C.112910). Fig. 5. Laevilitorina mariae: penis from adult ¢, Pirates Bay, Eaglehawk Neck, S. E. Tasmania, April, 1970 (Australian Museum C.112911). Figs. 6, 7. Laevilitorina johnstoni: shell of topotype, Ellensbrook, south Western Australia, 5.8 x 3.2 mm (Australian Museum C.114546). 778 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Plate 2. Figs. 1-4. Littorina (Austrolittorina) sundaica: shells, Pangandaran, Java, Indo- nesia (USNM 766856); figs. 1, 3, d 5 x 2.6 mm; figs. 2, 4, 9 5.6 x 2.9mm. Fig. 5, Littorina (Austrolittorina) acutispira: radula x 1300, Edwards Beach, Balmoral, Sydney, New South Wales, Australia (Australian Museum, C.112910). Fig. 6. Laevilitorina mariae: radula x 1400. Pirates Bay, Eaglehawk Neck, S. E. Tasmania, April 1970 (Australian Museum C.112911). species although the available samples are not large. One male was found among 9 specimens examined (detected by the presence of a penis; females by the absence of a penis). In general, apertures of females appear propor- tionally wider. Spiral sculpture appears more marked in the additional material now avail- able and ranges from a few spirally incised furrows most noticeable on the body whorl above the periphery, to a fairly uniform spirally furrowed con- VOLUME 92, NUMBER 4 779 dition (see Pl. 2, figs. 1-4). Other comments in the description made by Rosewater (ibid.) still apply. Laevilitorina johnstoni (Cotton, 1945) Plate wn, figs.:6,07 Pellax johnstoni Cotton, 1945:164, pl. 12, figs. 3, 4. Type: Holotype (South Australian Museum, D.14200). Ellenbrook [=Ellensbrook], south West- ern Australia. Laevilitorina burni Ponder, 1976:106, text-fig. 8; pl. 1, fig. 1; pl. 2, figs. 5, 6. Types: Holotype (Australian Museum, Sydney, C.100919, with 9 para- types; 3 paratypes Western Australian Museum). Yallingup, south West- ern Australia. Remarks.—This species was first introduced as a member of the Phasi- anellidae, presumably because of its striking color pattern. It closely resem- bles Eatoniella (Pellax) flammulata (Hutton, 1878), the type-species of Pel- lax Finlay, 1927 (=Phasianella huttoni Pilsbry, 1888, an unnecessary replacement name) and consequently was tentatively regarded as a member of the Eatoniellidae by one of us (W.F.P.) pending examination of the ani- mal. The radula and operculum of some half-grown specimens showing the characteristic color pattern of johnstoni proved to be identical to those of Laevilitorina burni Ponder, 1976. Comparison of all the material now avail- able has shown that L. burni and P. johnstoni are the same species and can be tentatively included in Laevilitorina Pfeffer, 1886, until its generic po- sition can be confirmed by the examination of its penis. This species appears to be confined to south Western Australia and South Australia. Laevilitorina mariae (T. Woods, 1876) Bleatongs. 5, Pl. 2, fis 6 Rissoa (Cingula) mariae T. Woods, 1876:147. Types: 4 syntypes (1 badly broken) (Tasmanian Museum, Hobart, TM7800/E459). King’s Island (Bass Strait, southern Australia). Remarks.—Ponder (1966) included this species in Rissolittorina Ponder, 1966 on shell characters, but examination of the animal has shown that it is best retained in Laevilitorina where it was placed by Hedley (1906). Ponder (1976) showed that the type-species of Laevilitorina, Littorina cal- iginosa Gould, 1849, has a simple penis whereas the penis of Rissolittorina alta (Powell, 1940) has a long accessory appendage (Ponder, 1966). The radula (Pl. 2, fig. 6) and penis (PI. 1, fig. 5) of L. mariae are illustrated. This Species is common in Tasmania and Victoria, extends into South Australia and is usually uncommon in New South Wales. 780 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Ml 0.5mm Plate 3. Figs. 1, 2. Littorina (Austrolittorina) sundaica: 1, Radula; 2, Penis, both taken from same male specimen (5 x 2.6 mm) from Pangandaran, Java, Indonesia (USNM 766856). Fig. 3. Littorina (Austrolittorina) praetermissa: penis from adult 6, Whisky Bay, S.W. side Wilson’s Promontory, Victoria, Australia, Dec. 1971 (Australian Museum C. 112912). Additional Notes Other changes in status of Indo-Pacific Littorinidae brought to our atten- tion since Rosewater’s (1970) review are: Littorina hisseyiana T. Woods, 1876: Rosewater (1970:476) correctly con- cluded that this species is a trochacean. It can be added that Kershaw VOLUME 92, NUMBER 4 781 (1955:291) has erected for it the genus Hisseyagibbula which he places in the Trochidae. Littorina incisa Yokoyama, 1927: Rosewater (1970:466) included this species in Littorinopsis, but according to T. Habe (in litt., 1971) it is a member of the Family Pyramidellidae. Littorina lucida Yokoyama, 1927: Rosewater (1970:453) included this species in the subgenus Littoraria. According to T. Habe (in litt., 1971) it is not a littorinid but is the species Assiminea japonica Martens, 1877. Acknowledgments Dr. T. Habe provided information on the status of L. incisa and L. lucida. Dr. C. Roper contacted A. Budiman for one of us (JR) who kindly provided the specimens of L. sundaica. We thank Mr. E. K. Yoo who prepared the radulae for S.E.M. examination and Ms. B. Duckworth and Mr. Yoo for assistance in preparing some of the line drawings and plates. Literature Cited Altena, C. O. van Regteren. 1945. Report upon a collection of Recent shells from Java.— Zoologische Mededeelingen, Leiden, 25:140-154. Cotton, B. C. 1945. Southern Australian Gastropoda. Part 1. Streptoneura.—Trans. R. Soc. S. Aust. 69(1): 150-171. Gray, J. E. 1826. In P. P. King, Narrative of a Survey of the Coasts of Australia, vol. 2, Appendix B, pp. 474-496. Hedley, C. 1906. Studies on Australian Mollusca. Part 9.—Proc. Linn. Soc. New South Wales 30:520—-546. . 1913. Studies on Australian Mollusca. Part 11.—Proc. Linn. Soc. New South Wales 38:258-339. Kershaw, R. C. 1955. A systematic list of the Mollusca of Tasmania, Australia.—Pap. Proc. Roy. Soc. Tasmania 89:289-355. May, W. L. 1909. Additions to the Tasmanian molluscan fauna.—Pap. Proc. Roy. Soc. Tas- mania for 1908:53-58, pl. 6. Macpherson, J. H. 1966. Brachiopoda and Mollusca.—Mem. Nat. Mus. Vict. 27:199-284. Macpherson, J. H. and C. J. Gabriel. 1962. Marine Mollusca of Victoria, pp. i-xv, 1-475.— Melbourne University Press. Ponder, W. F. 1966. The New Zealand species previously known as Zelaxitas Finlay, 1927 (Mollusca, Gastropoda).—Rec. Domin. Mus., Wellington 5(17): 163-176. . 1976. Three species of Littorinidae from southern Australia.—Malac. Rev. 9:105—114. Pritchard, G. B. and J. H. Gatliff. 1902. Catalogue of the Marine Shells of Victoria. Part 5.— Proc. Roy. Soc. Victoria (n.s.) 14(2):85—138. Reeve, L. A. 1857-58. Monograph of the Genus Littorina.—Conchologia Iconica, vol. 10, Littorina, plates 1-16 (1857), plates 17-18 (1858). Rosewater, J. 1970. The family Littorinidae in the Indo-Pacific. Part 1. The subfamily Litto- rininae.—Indo-Pacific Mollusca 2(11):417—5S06. 782 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Smith, E. A. 1892. Descriptions of new species of shells from New South Wales, New Guinea, the Caroline and Solomon Islands.—Proc. Zool. Soc. London for 1891:486—491. Woods, J. E. Tenison. 1876. Description of new Tasmanian shells.—Pap. Proc. Roy. Soc. Tasmania for 1875:134—162. . 1877. Census, with brief descriptions of the marine shells of Tasmania and the adjacent islands.—Pap. Proc. R. Soc. Tasmania for 1877:26—S7. (WFP) The Australian Museum, 6-8 College Street, Sydney 2000, Aus- tralia; (JR) National Museum of Natural History, Washington, D.C., 20560. PROC. BIOL. SOC. WASH. 92(4), 1979, pp. 783-795 SEX-REVERSAL AND VIVIPARITY IN OPHIOLEPIS KIERI, N. SP., WITH NOTES ON VIVIPAROUS BRITTLESTARS FROM THE CARIBBEAN (ECHINODERMATA: OPHIUROIDEA) Gordon Hendler Abstract.—A new species, Ophiolepis kieri, appears to be more closely allied to Ophiolepis impressa than to its other Caribbean congeners Ophi- olepis elegans and Ophiolepis paucispina. Three sympatric species Ophi- olepis kieri, Ophiolepis paucispina, and Ophiomyxa flaccida brood their young, bringing the number of known viviparous, tropical shallow-water ophiuroid species to seven. Ophiolepis kieri probably broods sequentially, but O. paucispina carries more than one clutch simultaneously. Ophiomyxa flaccida is a gonochore, O. paucispina is a simultaneous hermaphrodite, and O. kieri is a protandric hermaphrodite. A speculative sequence is pre- sented for the rapid evolution of small Ophiolepis species through paedo- morphosis, facilitated by preadaptations for viviparity and selection for her- maphroditism. The 57 species of viviparous ophiuroids that have been described are predominantly coldwater or deepwater forms (Bernasconi, 1965; Madsen, 1967; Mortensen, 1936; Murakami, 1941). Livebearers from shallow, trop- ical waters are so rare that Mortensen (1933) remarked on the novelty of the only examples: Amphiura stimpsoni Lutken, Caribbean; Cryptopelta granulifera H. L. Clark, Mascarene Islands and Northern Australia; Ophi- oconis vivipara Mortensen, Northwest Africa; and Ophionereis vivipara Mortensen, Mozambique and Mascarene Islands. Three additional species of tropical viviparous ophiuroids collected from the Caribbean coast of Panama are discussed below. Two of them have widespread distributions: Ophiomyxa flaccida (Say) ranges from Bermuda to Brazil and Ophiolepis paucispina (Say) from Bermuda to Brazil and West Africa (H. L. Clark, 1919; Madsen, 1970). The third species, described in this paper as Ophiolepis kieri, has been collected only at Portobelo, Panama (9°33'30"N, 79°40'45’W). It is the fourth Ophiolepis reported from the West- ern Atlantic and is sympatric with Ophiolepis impressa Lutken and Ophi- olepis paucispina (Say). Dissections that demonstrated brooding in these species also revealed several different types of sexuality. Specimens of O. flaccida proved to be gonochoristic, like most ophiuroids, and specimens of O. paucispina were 784 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON simultaneous hermaphrodites as are a majority of the viviparous brittlestars. However, the protandric hermaphroditism found in O. kieri was an excep- tional discovery. Although protandry had previously been reported in ophiuroids, the pro- gressive stages of sex-reversal involving small males, hermaphrodites of intermediate size, and large females were documented convincingly for only one species, Amphiura stepanovii Tscherniawsky (Fedotov, 1926). There- fore, the process of sexual transformation in O. kieri is described in detail below, and other notable aspects of the species are discussed. Ophiolepis kieri, new species Fig. 1 Etymology.—Named for Porter M. Kier, an echinodermologist of out- standing stature, to honor his retirement as Director of the National Museum of Natural History. Material examined.—Holotype (USNM E18177) and 18 paratypes (USNM E18175) collected 2 September 1977, Iron Castle Point, Portobelo, Panama, on sand under rocks, less than 2 meters depth, by the author. Ten paratypes (USNM E18176) collected 15 October 1975, Iron Castle Point, Portobelo, Panama, under rocks, less than 4 meters depth, by the author. Description of holotype.—Disc diameter 4.7 mm; longest arm 11.9 mm, all arms broken before tips. Disc circular, slight indentations at bases of arms; dorsal side convex; ventral side flat. Arms gradually tapering. Major disc scales with lateral and distal edges thickened; bordered but usually not completely surrounded by small scales. Primary plates forming distinct ro- sette; central and radial plates larger than any dorsal scales but radial shields. Radial plates ovoid; middle of long distal edge straight. Interradial field of disc with 3 columns of major scales; 3 scales in middle column larger than scales in lateral columns. Proximal scale in middle col- umn as long as wide, central scale longer than wide, peripheral scale wider than long. Radial field of disc with: 3 major scales distal to each radial plate; elongate scale between radial shields; supplementary radial shields linked by small scale, paired supplementary radial shields and connecting scales forming crescentric shape. Surface of some radial shields minutely wrinkled. Each jaw bearing pointed apical papilla plus 5 pairs lateral papillae. Prox- imal pair robust, pyramidal; next pair smaller, flat, rhomboidal, proximal edge sometimes drawn to point; succeeding pair similar but lacking acute point; next pair much larger, shaped like quarter-circle, radial edge curved; last pair long, triangular, attached to adoral shield, tip extending beneath penultimate pair. Oral shield slightly longer than wide; proximal angle acute, margins con- cave; lateral margins nearly parallel; distal margin deeply convex. Madre- VOLUME 92, NUMBER 4 785 Fig. 1. Ophiolepis kieri, holotype. A, Dorsal view of disc and of arm; B, Ventral view of two jaws with contiguous arm and interradial fields of disc; C, Lateral view of basal arm segments and part of adjoining disc. Scale lines equal 1 mm. porite resembling oral shield, larger; with small depression near center. Adoral shield widest distally; radial edge straight, adradial edge indented at point of contact with oral shield; surface of shield inflated. Genital slit extending nearly to edge of disc; bordered by long genital plate, 2 smaller scales proximad. Ventral interbrachial field with fewer than 25 scales. Dorsal arm plates triangular; basal plates with concave lateral edges, dis- tal edge with straight medial margin; remaining dorsal arm plates with straight lateral edges, distal edge with small blunt median projection. First 10 to 11 dorsal arm plates overlapping, remaining dorsal arm plates sepa- rated by lateral arm plates. Supplementary dorsal arm plates present only on first 7 to 10 arm seg- ments; miniscule, all except first pair smaller than arm spine. Lateral arm plates flaring in dorsal view; trapezoidal, widening distally in lateral aspect. Arm spines generally aligned with arm, not erect; tips of spines attenuate, middle thick. Ventralmost spine longest until segment 13 to 19, then same size as dorsal spines. Spines equal or less than length of ventral arm plate; spines on third segment beyond disc characteristically shorter than ventral arm plate. Spines often lacking on first lateral arm plate; increasing in num- ber to 3 per lateral arm plate, then decreasing to 2 as indicated in Table 1. 786 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON a NaN of ~~ i ANN xT | SS o N 2 aan Y a N Q NAN x N x ANNAN N Q Anan N oa) N man = N ing) a (oa) a NAANMAAM no SIP MAmManmnmnnnm ah, na (oo) ay ANNA NNANANM ae es AE Gal GN) SIE MMNnananaanm = ZiIet amas ol SPE mamnmnnnnnn = n 3 P ee) |} Sa NAAN no} No} 2) e| & ANAANANMNNM rf) 2 LN x NAAN (>) D Sh AOAANNAAMN NM a ns : 5 s ao N 2. 5 wt > BS ANNAN NAN MN M| P= SS) RED = ao} = TIT MNNMNMAANANAMNM 5 = An AN = ~~ o v AL nnanannan Ss =) 1) 0S? ‘SOE SCOR ~ ~ a is a analnn ann 5. me ANA AANANMN NM = : S 3 = 00 an AM is = NAA ANANANANANAMNM 3S & & ~t am am eo, EN AAA ANNA NANM OM bs le) io) Ne} ana mo N Dn 2 20 NAANANANAAMNN MN oO = a mt an mo a a i oa) 5 NANAMNMNANNM - a Poe ares oc a ke a Ne} NANA AANA MN MH a o|™ man In o-m (o) (e) we Z Es n < 5 Vay ANAMAANAMNMNM NM c == Trica of bee, 4 2 |a8 3 +t NAAM AMNNNMNM MN = rile ~T e ie (=| Gey ° ro IM|ENANANANANANANAN ° ayejd we 72) 2 25 e jeiajey 13d =) oO : c Be = jo Joquinny fe) |) o 7p) Siete Cle ole oS — nH =n +0 | | Sans = N ) = a Es} d 9)e] UO = es 9}e]| wie [el Je] SS JIg}oweIp souids wie Jo 19quinyy osiq wu VOLUME 92, NUMBER 4 787 Ventral arm plates beyond segments 7 to 9 separated by lateral arm plates; flat, bell-shaped, with proximal angle acute, lateral margins concave, distal margin convex, distal end with pointed lateral lobes. Basal ventral arm plates overlapping, proximal angle concealed, surface of plate concave. Plates beneath disc with straight mid-distal margin. Two rounded triangular tentacle scales on each side of arm until segment 20 to 21. Paired scales over each pore forming an oval operculum. Long axis of opercula on proximal segments half or less than half length of ventral arm plate; length of operculum on distal segments equal or exceeding length of ventral arm plate; length of operculum on third segment beyond disc always less than half length of ventral arm plate. Minute, supplementary tentacle scales on some basal segments. Color (in alcohol): Disc above mottled pale pink-gray, some major scales tipped with white or brown, some small scales white; disc below white. Arms above lighter than disc, with iregular blotches or bands; arms below white. Dorsal arm plates mottled with pale brown and gray, some lighter in center than along margins. Lateral arm plates lighter than dorsal arm plates. Variations.—Observations below are based on the paratype series. Dimensions: For specimens 1.9 to 5.5 mm disc diameter y = 3.41x — 3.37, where y = arm length and x = disc diameter (n = 27, r = 0.95). Disc scalation: The prominence and shape of the primary (central and radial) plates is consistent throughout a size series from 1.9 to 5.5 mm d.d. The number of ventral interbrachial scales varies directly with size, a spec- imen 5.5 mm d.d. having but a few more scales than the holotype. Speci- mens 2.6 mm d.d. and less have a reduced number of small scales and lack scales between the supplementary radial shields. Specimens less than 2.0 mm d.d. have the radial shields in direct contact and their major scales are reduced in number. Arm plates: With decreasing body size there is less overlapping both of adjacent dorsal arm plates and of adjacent ventral arm plates. In the smallest specimens, dorsal and ventral arm plates are both separated by the lateral arm plates for nearly the full length of the arm. The first arm segment with the dorsal arm plate surrounded by lateral arm plates is, in a specimen: 3.4 mm d.d., at segment 4 beyond the disc; 3.2 mm d.d., segment 3; 2.6 mm d.d., segment 2; 1.9 mm d.d., segment |. The first arm segment with the ventral arm plate surrounded by lateral arm plates is, in a specimen: 3.7 mm d.d., at segment 6; 3.4 mm d.d., segment 4; 3.2 mm d.d., segment 4; 2.6 mm d.d., segment 3; 1.9 mm d.d., segment 2. The distal edge of the ventral arm plate is rounded rather than pointed in small specimens. Concavity of the surface of the ventral arm plates is not pronounced in small specimens, but is still perceptible in an ophiuroid 1.9 mm d.d. The number of arm segments beneath the disc varies directly with body size: 3 segments are covered in specimens 4.1 mm d.d. or larger, and 2 segments are covered in specimens 3.6 mm d.d. or smaller. 788 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Arm spines: The absence of spines on the first lateral arm plates is most pronounced in larger specimens, indicating that some spines are resorbed as individuals mature. Specimens at least 3.4 mm d.d. average fewer than 5 first lateral arm plates carrying spines, and for specimens less than 3.2 mm d.d. an average of fewer than 7 first lateral arm plates bear spines. For specimens of different sizes, the number of arm spines along one side of a typical, unbroken arm varies as indicated in Table 2. Tentacle scales: Scales over the tentacle pores are paired. Distally the radial scale of the pair becomes increasingly smaller than the abradial scale and disappears near the arm tip. Smaller specimens have fewer segments with paired tentacle scales. The first segment with only one scale over each pore is, for a specimen: 5.5 mm d.d., segment 26; 4.1 mm d.d., segment 20; 3.7 mm d.d., segment 18; 3.4 mm d.d., segment 14; 3.2 mm d.d., segment 13; 2.6 mm d.d., segment 8; 1.9 mm d.d., segment 6. Color (in alcohol): Specimens less than 3.2 mm d.d. are pale beige; those less than 2.9 mm d.d. are entirely white. Relationships.—Ophiolepis kieri differs dramatically from congeners such as O. elegans Liitken and O. paucispina (Say) which, even as juveniles, have a row of small scales of uniform size surrounding the major disc scales and have large supplementary dorsal arm plates on most arm segments. Ophiolepis kieri, with incomplete, irregular rows of small disc scales and with supplementary dorsal arm plates of reduced size and number is more Closely allied to Ophiolepis impressa Liitken and Ophiolepis pacifica Lut- ken. The latter species are relatively large, exceeding the size of the biggest (6.4 mm d.d.) O. kieri. However, there is a striking similarity in the ontog- eny of disc scales of the three species, the disc armament of O. kieri resem- bling that of young O. pacifica and O. impressa. Ophiolepis kieri may be distinguished from small O. pacifica and O. impressa as follows: O. kieri has 3 rather than 4 or more arm spines on the basal arm segments; 3 rather than 4 or more major scales in the central scale column of the dorsal inter- radial field; primary radial plates that form a ring, and are larger than other dorsal scales except the radial shields. Specimens of O. impressa also differ from O. kieri because their dorsal arm plates are in contact for over half the length of the arm, their first lateral arm plates each bear 2 spines, and the length of their tentacle-scale pairs is generally greater than half the length of the ventral arm plate. Ecology.—Ophiolepis kieri has been collected only at Portobelo, Panama, in the lee of Iron Castle Point. The species inhabits a shallow, sheltered sublittoral zone where numerous allochthanous boulders and cobbles pave a substratum of fine sand. The rocks bear encrusting calcareous algae, po- rifera, bryozoa, and Millepora. Ophiolepis kieri is found only up to 3 m. At VOLUME 92, NUMBER 4 789 that depth, the rocky habitat abuts a steep reef slope supporting large corals and sponges. The reef ends at 20 m on a calcareous mud bottom. Salinities at Iron Castle Point ranged from 32 to 37%oc but it is likely that the very heavy rains in this locality create temporary, steep salinity and temperature gradients in shallow water. Under the rocks at the type-locality there were dense populations of ophiuroids including, in order of abundance: Ophiocoma echinata, Ophi- ocoma wendti, Ophiothrix angulata, Ophioderma rubicundum, Ophiocoma pumila, Ophioderma appressum, as well as rarer species of ophiuroids and conspicuous macroinvertebrates such as brachyura, actiniaria, and poly- chaeta. Specimens of Ophiolepis kieri were found only beneath cobbles firmly lodged in fine, dark sand. Under any particular rock they were either found in numbers or were absent, but they were never found under rocks shelter- ing macroinvertebrates other than their congeners, O. impressa and O. paucispina. Ophiolepis kieri was more numerous than O. paucispina. In a series of collections at Portobelo, the ratio of the two species was 103:30. When their rock shelters were dislodged, the three sympatric Ophiolepis species ‘‘froze’’ rather than moving to escape, and their variegated, dusky coloration camouflaged their location. Labrid fish feed on ophiuroids at Portobelo, but the protected microhabitat and behavior of O. kieri probably reduce predation mortality. The same factors also minimize the physical disturbance of these lilliputian forms by large active organisms. Notes on viviparous Caribbean Ophiuroids Ophiolepis kieri.—Male, female, and hermaphrodite specimens differed in body size (Fig. 2). Males ranged from 1.9 to 5.0 mm d.d. (¢ = 3.6 mm). They had a single, white testis near each genital slit. The testes contained spermatozoa with tails at least 10 times the length of the spherical sper- matozoan head. The transitional, hermaphroditic individuals were intermediate in size be- tween males and females (3.8 to 5.1 mm d.d., x = 4.5 mm). Their gonads were pure testes, pure Ovaries, or ovotestes. Hermaphroditic individuals might functions as males, but not as females. Their ovaries and ovotestes contained small, incompletely yolked oocytes and they did not brood em- bryos. Although some ovaries may form de novo in the hermaphrodites, it is obvious that testes transform to ovotestes, and the ovotestes become Ovaries. Females were 4.5 to 6.4 mm d.d. (¢ = 5.0 mm) and had 1-2 ovaries with grayish-violet oocytes, at each genital slit. The yolky, opaque oocytes reached a maximum diameter of 0.35 mm. The maximum number of fully 790 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON developed oocytes in an individual was 32, and the maximum number of embryos was 39 (n = 13). The embryos in a female were generally all at the same stage of development; usually either early stages, small stars with terminal arm plates, or juveniles with 1 to 2 arm segments. Brooding females, and males with active spermatozoa, were collected in January, February, March, April, June, August, and December, indicating there might be a long breeding season and recruitment period. Newly hatched young were rarely encountered. The size-frequency curve (Fig. 2) shows the distribution of males to be skewed toward the larger sized indi- viduals, a scarcity of transitional hermaphroditic forms, and a restricted, unimodal distribution for females. The size-frequency curve suggests delayed sex-reversal of the males. As- suming the population is in equilibrium and that size-specific mortality of males is not significant, there appears to be infrequent, but rapid, transfor- mation from males to females. It is possible that protandry is labile and that sexual transformation is retarded in the presence of mature females. However, the pronounced seg- regation by size of males and females may indicate that sex-reversal is genetically predetermined rather than environmentally controlled. Whatever the cause, the bottleneck to growth or sex-reversal contributes to mainte- nance of a population dominated by males. The sex ratio is 2.4:1 (hypothesis that the ratio is 1:1 is rejected: x?, P < 0.005). Such a proportion would tend to ensure that females would be fertilized. Ophiolepis paucispina.—Nearly all individuals from Portobelo were si- multaneous hermaphrodites. Those from 3.2 to 5.4 mm d.d. had functional testes, and ovaries with large oocytes. There was no tendency for the larger individuals to have a greater number of ovaries (and fewer testes) and thus no indication of protandry (hypothesis that the Kendall coefficient of rank correlation equals zero not rejected, P > 0.99). Only 3 of the 30 O. pau- cispina dissected were males, and these were large (6.2 mm d.d.) as well as small (4.0 mm d.d.) individuals. There was generally one gonad to each genital slit, and the testes and Ovaries were ordered in an irregular sequence around the disc. However, series of consecutive genital slits with only male or female gonads some- times formed statistically significant ‘‘runs.’’ There was a ‘‘functional sex- ratio’ favoring females since 170 ovaries, but only 80 testes and 2 ovotestes were counted in 30 individuals. The largest oocytes were 0.35 mm in diameter, opaque and pink. The maximum, number of fully developed oocytes found in an individual was 16, and the maximum number of embryos was 41 (n = 11). Specimens brooding young were collected in March, April, June, and December and individuals often contained embryos at different stages of development. Broods consisted of yolky embryos in early developmental stages or juve- VOLUME 92, NUMBER 4 791 25 Ophiolepis kieri cere MALE (——] HERMAPHRODITE INDETERMINATE 20 N = 103 15 10 NUMBER OF SPECIMENS 1.6 3.0 3.5 4.0 4.5 50 54 5.9 6.4 DISC DIAMETER (mm) Fig. 2. -Size-frequency histogram for the population of Ophiolepis kieri sampled at Porto- belo, Panama, showing the transformation during maturation through male, hermaphrodite and female phases. Indeterminate individuals lacked gonads with recognizable gametes. 22.6 niles with 2 to 3 arm segments. White testes with active spermatozoa were found both in brooding individuals and those not brooding embryos. Although O. kieri and O. paucispina brood approximately equal numbers of embryos, individuals of the latter species produce a smaller “‘clutch”’ of eggs and brood embryos of different ages within the same bursa. This sug- gests that O. kieri is a sequential brooder but O. paucispina simultaneously broods embryos from more than one clutch of eggs. Viviparous ophiuroids such as Axiognathus squamata, which have ova- ries and testes associated with every bursa, may bear embryos simulta- neously in all their bursae (personal observation). In contrast, in O. pau- cispina all of the bursae next to testes were empty, while 59% of the bursae associated with ovaries contained embryos. It would be easier to interpret this phenomenon if the position of the gonoduct in relation to the bursa 792 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON were known, but it is likely that after spawning the eggs are retained in bursae into which they are shed. Because ‘‘male bursae’’ apparently cannot brood, the uneven ‘‘functional sex ratio’? mentioned above may be an ad- aptation to increase the number of brood-bearing bursae and to increase fecundity. The ‘‘division of labor’’ between male and female bursae might function to prevent self-fertilization, and also may figure in the mechanism coordinating the number of oocytes released by an ovary, with the number of embryos that are brooded in the associated bursa. Ophiomyxa flaccida.—Fifty-two specimens were collected at Portobelo and approximately 25 km away at Galeta Reef (9°24’18"N, 79°51'48.5”W) Both populations were strictly gonochoristic. Males were slightly smaller than females (mean disc diameters 1.72 vs. 1.93 mm), and considerably less common. Only 32% of the individuals examined were male (hypothesis that the sex ratio is 1:1 rejected, x?, P < 0.005). The coloration of individuals varied from green to red; some specimens had uniform coloration and others were mottled with patches of intense pigmentation. Brooding females were found only at Galeta Reef. However, the absence of brooding individuals from Portobelo may be a sampling artifact, as only 10 females were collected at that site. The brooding females each held up to 48 embryos and the embryos in a female were generally at the same stage of development; in only one case were early and advanced embryos found together in an individual. The largest embryos had up to 6 arm segments. Brooding females were collected in March, May, June and July, but all females collected in November, December, and February were without young. Ripe testes, with active long-tailed spermatozoa, were found in males throughout the year. Unfortunately, too few brooding specimens were found to detect reproductive periodicity. Egg counts and measurements, though taken only for a few specimens, suggest important differences between brooding females and those without young. The largest oocytes were roughly the same diameter in brooding females (0.75 mm, n = 2 specimens) and females lacking embryos (0.70 to 0.88 mm, n = 3 specimens). In brooders, as well as females without young, the immature oocytes were translucent pinkish-gray and the yolky oocytes were Opaque green. However, ripe oocytes teased from the gonads of non- brooding females were buoyant but those from brooding females were heavi- er than seawater. Furthermore, brooding females (1.76 and 1.82 cm d.d.) carried approximately 260 and 200 ripe oocytes respectively, while a non- brooding individual (1.95 cm d.d.) had approximately 915 oocytes. Changes in yolk composition of oocytes before shedding and lowered oocyte counts due to spawning, could account for the variability of the oocytes noted above. However, a significant difference between the mean disc diameter of brooding females (1.52 cm, n = 7) and of those without broods (2.05 cm, n = 24) (Wilcoxon two-sample test, 0.02 > P > 0.01), sug-: VOLUME 92, NUMBER 4 oo gests the information on oocytes figures in an alternative explanation for the contrasts between brooding and “‘barren’’ females. Namely, it is possible that small female Ophiomyxa flaccida have small numbers of dense eggs and brooded embryos, but larger females produce many floating eggs and planktonic larvae. Obviously, additional observations are needed to sub- stantiate this hypothesis, and a planktonic Ophiomyxa larva, if one exists, remains to be discovered. Discussion Information on the reproductive biology and morphology of the Pana- manian Ophiolepis species suggests a hypothetical sequence for the evo- lution of some members of the genus. The smallest Ophiolepis species may simply have originated from selection for reduced size. On the other hand, the diminutive species might have evolved from paedogenetic forms with early sexual maturity, “‘juvenile’’ somatic development; and with their suc- cess enhanced by viviparity and hermaphroditism. Two groups of Panamanian Ophiolepis species may be distinguished by differences in their disc scalation. One set is composed of O. kieri, O. pacifica and O. impressa, which have an irregular or incomplete row of small scales around the major dorsal disc scales, and have inconspicuous supplementary dorsal arm plates. Another set, composed of O. paucispina, O. elegans, and O. variegata have a uniform border of small scales around each of the major dorsal disc scales, and have large supplementary dorsal arm plates. Further systematic work is needed to determine whether these two groups are valid natural assemblages. While certain specialized features, such as the arrangement of the oral papillae, exhibit the same degree of complexity in the small and large rep- resentatives of both species sets, there are other features, such as disc scalation that are simplified in the smaller species in both groups. The pat- tern of disc scalation increases in complexity during ontogeny, as well as according to size within a series of allied species. For example, the scale pattern of a small form such as O. kieri approximates that of juveniles of larger species such as O. impressa and O. pacifica. The co-occurrence of simplified somatic growth and sexual maturity at a small size in the diminutive Ophiolepis species, O. kieri and O. paucispina, suggests that they evolved by paedogenesis (progenesis, sensu Gould, 1977). If this were so and a tendency to produce yolky eggs of moderate size were fixed in the genotype of the small species, their potential fecun- dity, limited by the room for oocytes in the disc, would have been consid- erably lower than that of their larger congeners. In this situation, the dis- advantages of low fecundity would have been mitigated by an adaptation such as brooding, that enhanced the survival of young. 794 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON There is a tendency for both large and small Ophiolepis species to produce yolky eggs of moderate size and modified embryos or larvae. Ophiolepis kieri and O. paucispina have yolky eggs and are viviparous, while O. ele- gans and O. cincta have yolky eggs, lecithotrophic larvae, and abbreviated development (Mortensen, 1938; Stancyk, 1973). It is likely that a Caribbean species, Ophiolepis impressa, and its Eastern Pacific congener, Ophiolepis pacifica, also have abbreviated development. Females of both species do not brood, but have dark brown, opaque yolky oocytes 0.2 mm in diameter (personal observation). Eggs of that size typically undergo abbreviated de- velopment (Hendler, 1975). The yolky eggs and abbreviated development characteristic of the genus could be preadaptations for viviparity. Consequently, in the genus Ophiolepis, paedogenesis coupled with pre- adaptations such as yolky eggs, modified development, and genital bursae (brood sacs), in concert with selective pressure for brooding because of low fecundity, may have contributed to rapid evolution of small size and vivi- parity. The chief evidence for paedogenesis, the juvenile (not simply minia- ture) appearance of the small species, does not exclude the alternative pos- sibility of gradual selection for small body size. However, evidence for the speculative sequence of events above might be found through further study of the reproduction, ontogeny, and phylogenetic relationships of the genus Ophiolepis, and by inference from similar trends for paedogenesis associ- ated with abbreviated development and viviparity in other genera. The significance of hermaphroditism in viviparous species of Ophiolepis, in fact the reason that all hermaphroditic ophiuroids are viviparous, is as yet unexplained (Hendler, 1975). The most appropriate hypotheses are the ‘“‘density model’’ which suggests that hermaphroditism increases the like- lihood for sedentary organisms to encounter a mate, and the “‘resource allocation model’’ which suggests that hermaphroditism increases the effi- ciency of apportioning energy between male and female functions (Smith, 1978). Either of these hypotheses helps explain the adaptive importance of hermaphroditism in these relatively small, rare brooding brittlestars. Acknowledgments I tender special thanks to Janice Clark for her illustration of Ophiolepis kieri and to Maureen Downey, David Franz, David Pawson, and Lowell Thomas for their comments on the manuscript. In addition, support from the Smithsonian Institution through a Walter Rathbone Bacon Scholarship and the Secretary’s Fluid Research Fund is noted with gratitude. Literature Cited Bernasconi, I. 1965. Astrotoma agassizii Lyman, especie vivipara del Atlantico Sur (Ophiu- roidea, Gorgonocephalidae).—Physis 25:1—S. VOLUME 92, NUMBER 4 1D Clark, H. L. 1919. The distribution of the littoral Echinoderms of the West Indies.—Carnegie Inst. Publ. No. 281:49-74. Fedotov, D. 1926. Zur Morphologie einiger typischen, vorzugsweise lebendiggebarenden, Ophiuren.—Trav. Lab. Zool. Stat. Biol. Sevastopol Ser. 2, 6:39-72. Gould, S. J. 1977. Ontogeny and Phylogeny.—Harvard University Press. Cambridge, Mas- sachusetts. 501 pp. Hendler, G. 1975. Adaptational significance of the patterns of Ophiuroid development.— Amer. Zool. 15:691-715. Madsen, F. J. 1967. Ophiuroidea.—B.A.N.Z. Antarctic Research Expedition, 1929-1931. Se- ries B. 9:121-145. . 1970. West African Ophiuroids.—Atlantide Report No. 11:151-243. Mortensen, T. 1933. Biological observations on Ophiurids, with descriptions of two new genera and four new species.—Vidensk. Medd. Naturhist. Foren. Kgbenhavn 93:171- 194. . 1936. Echinoidea and Ophiuroidea.—Discovery Reports. 12:199-348. . 1938. Contributions to the study of the development and larval forms of Echinoderms IV.—D. Kgl. Dansk Vidensk. Selsk. Skr. Naturv. Math. 7:1-5S9. Murakami, S. 1941. On the development of the hard parts of a viviparous Ophiuran, Stego- phiura sculpta (Duncan).—Annot. Zool. Japon. 20:67-78. Smith, J. M. 1978. The Evolution of Sex.—Cambridge University Press. Cambridge, England. x 4 222 pp: Stancyk, S. E. 1973. Development of Ophiolepis elegans (Echinodermata: Ophiuroidea) and its implications in the estuarine environment.—Mar. Biol. 21:7—12. Smithsonian Oceanographic Sorting Center, Smithsonian Institution, Washington, D.C. 20560. PROC. BIOL. SOC. WASH. 92(4), 1979, pp. 796-800 CLYPEASTER KIERI, A NEW SPECIES OF CLYPEASTEROID (ECHINODERMATA: ECHINOIDEA) FROM OFF BOMBAY, INDIA David L. Pawson and Thomas F. Phelan Abstract.—A new species of the clypeasteroid echinoid genus Clypeaster is described from 96-106 m depth off Bombay, India. The new species, C. kieri, falls within the ‘‘Stolonoclypeus”’ section of the genus, where it ap- pears to have no close relatives. Among the large collections of echinoids amassed by the International Indian Ocean Expedition and entrusted to D.L.P. for study are specimens of a new species of Clypeaster. The new species is described here. Type- material is deposited in the National Museum of Natural History, Smith- sonian Institution (USNM). Class Echinoidea Family Clypeasteridae L. Agassiz, 1835 Clypeaster Lamarck, 1801 Clypeaster kieri, new species Diagnosis.—Test with flattened margin; central part raised. Marginal in- ternal skeleton well developed. Paired petals closed; anterior petal usually slightly open. Petaloid area 63-67% of test length. Ridges between pore- pairs of petals with 4—5S scattered tubercles. Material examined.—HOLOTYPE (USNM E18188), International Indian Ocean Expedition, Anton Bruun Cruise 4B, Sta. 202A, 17°25’N, 71°39’E to 17°21'N, 71°41'E, 13 November 1963, 96-106 m, greenish sand and mud. PARATYPES: 14 specimens (USNM E18189), same locality data as holo- type; 12 specimens (USNM E18190), Sta. 202B, 17°41’N, 71°33°E to 17°45'N, 71°32'’E, 14 November 1963, 90 m; 5 specimens (USNM E18191), Sta. 202C, 18°27'N, 71°13’E to 18°38’N, 71°09’E, 14 November 1963, 84—97 m, coral and sand. Etymology.—This species is named in honor of our colleague and friend Dr. Porter M. Kier, Director, National Museum of Natural History, Smith- sonian Institution. Description.—Holotype 50 mm long, 45 mm wide. Paratypes 16.1 to 55 mm long. Test longer than broad, length/width ratio ranging from 0.98 to 1.05 (see Table 1; all but one 42.8 mm long specimen longer than broad). “97 VOLUME 92, NUMBER 4 “ape j +i il ‘ ‘° 2. 2, aay 00 So. o a oo 9°o * ears &, °, 1% oe 5 vo Vv 22 aA or | (e) (©) fay a So wo « Bie > 2 a 6 royce ae > a e< os _d) Xoo ~ aboral v ] structure of 53 mm Paratype from Stat d central lamellae Clypeaster k spines remove Fig. 1. inal and ll devel- ll developed marg ing we 202A, show F, apical system of 33 mm Paratype from Stat oped genital pores. 10n interna ing we 202C, show 10n . 9 798 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Edge of test slightly indented in each interambulacrum, except that test edge is usually convex in posterior interambulacrum. Test relatively low, more or less flat above ambitus, then gradually rising to apex. Height 22.2—27.9% of test length. Edge of test thin, approximately 7% of test length. Mouth sunken; outer edges of oral surface essentially flat. Petaloid area occupies approximately 63-67% of test length. Paired petals closed distally; anterior unpaired petal usually open, with extremities of petal edges curving towards each other. In few specimens anterior petal completely closed. Anterior petal longest, posterolateral petals next longest, anterolateral petals shortest (see Table 1). In a 19 mm long anterior petal there are 30 pairs of pores; in an anterior petal 10 mm long there are 26 pairs. On ridges between pore- pairs, number of primary tubercles increases as animals grow; in largest specimens there are 4—5 tubercles per ridge and in small specimens there are 1-3. Tubercles always scattered, never closely aggregated. In interambulacra, primary tubercles evenly distributed; aborally there are 240-270 tubercles per cm?. Apical system slightly posterior; distance from anterior ambitus to apex is 54-56% of test length. Five genital pores do not appear until test length of 30 mm is attained. Peristome approximately opposite apical system, circular to laterally elongate, depressed. Periproct round to slightly irregular, lying no more than its own diameter anterior to posterior edge of test. Primary oral spines approximately 1.7 mm long, mostly cylindrical, straight to slightly curved. Primary aboral spines approximately 0.8 mm long, straight, gently tapering. Oral miliary spines straight, gently tapering, usually slightly widened at tips, 0.45—0.5 mm long; aboral miliaries similar, approximately 0.4 mm long. Tridentate pedicellariae typical, common, larger on the oral surface (length of valves averages 0.5 mm) than on aboral surface (valves 0.31 mm). Ophicephalous pedicellariae not common, occurring apparently only near ambitus; valves approximately 0.2 mm long. No other types of pedicellariae found. Internal skeleton consists of marginal lamellae in approximately 3 bands, forming short walls, and inner series of well developed and numerous la- mellae. Two lateral water vessels lie on inner surface of each primary petal plate, each serving a respiratory tube foot and accessory tube feet. On inner surface of adoral plates lateral water vessels are unbranched, lying almost at right angles to radial water vessels. Accessory pores lie to sides of lateral water vessels, towards adapical sutures of their respective plates. Remarks.—According to the various figures given by Mortensen (1948) and Serafy (1970, 1971), it would appear that the average size at which genital pores appear in young specimens of Clypeaster is approximately 35% of the maximum size attained by mature specimens. This average figure is based upon data from 16 species. As genital pores in C. kieri arise at a 799 VOLUME 92, NUMBER 4 ‘WNJoR[Nquiesoyut dulpuodsaiiod Ul snyiquie 0} xede woJy sduR}sSIP Jo adeJUI.10d & se possoidxa si yIBua] [ejod ‘oudH ,, “SIOJOWNI[IW UI oue s}USWOINseaW IV nn SSS 6° C/S9'7 LL’€/60°7 91 °€/7S 7 66 87 vy Le Ol cL'9 ‘a's 6 89/L'ST 1°S$9/8°€1 99/1 (GA\KG £0'l 6£ oe ueouwl CHL—S COV CHT CL €CL-E'SS/L LI-V II 8 IZ-1°6S/L'07-L IT Gee GGG S0' 1-86'0 CES-IE CS-P'0E osuel a eee % Se/yIBu9] «% Se/IBUD 4% Se/yyBug] TJowse A/T (A) (1) III [eed II [eted I [ried 1YsI0H oney UIPIAA yysueT eB ea ee ee ‘Maly dajspadaj) 10} eyep [eoNsnejs— | 21qQe], 800 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON test length of approximately 30 mm, it seems likely that this species attains a maximum length of approximately 85 mm. | The pattern of distribution of the water vessels and their branches con- forms to that described by Phelan (1977). This species falls within the ‘“Stolonoclypeus’’ section of the genus, stated by Mortensen (1948) to include species which have a test with flattened margin, with oral surface flattened or concave, with petals more or less distinctly closed, and with marginal internal skeleton well developed. Within this section of the genus, C. kieri appears to have no very close relatives, although some features are shared with the common Indo-west-Pacific species C. humilis (Leske). Conspicuous differences between these 2 species are: in C. humilis the petaloid area occupies little more than half the test length while in C. kieri it occupies 63-67% of test length. In C. humilis the anterior unpaired petal is usually closed, while in C. kieri it is usually open. In C. humilis there are dense regular series of tubercles in the ridges between the pore-pairs in the petals; small specimens (20-50 mm?) have 3—5 tubercles and larger specimens have up to 10. In C. kieri these tubercles are irregularly scattered, and there are 4—5 tubercles per ridge in larger specimens. In C. humilis genital pores do not appear until a length of 46-50 mm is attained, while in C. kieri they appear at a length of ap- proximately 30 mm. Literature Cited Mortensen, F. 1948. Clypeastroida, Clypeastridae, Arachnoididae, Fibulariidae, Laganidae and Scutellidae. Number 2 of Volume 4 in Mortensen.—A Monograph of the Echinoidea. 471 pages, 258 figures, 72 plates. Phelan, T. F. 1977. Comments on the water vascular system, food grooves, and ancestry of the clypeasteroid echinoids.—Bulletin of Marine Science 27(3):400-—422, 29 figures. Serafy, D. K. 1970. A new species of Clypeaster from the Gulf and Caribbean and a key to the species in the tropical northwestern Atlantic (Echinodermata: Echinoidea).—Bul- letin of Marine Science 20(3):662—677, 7 figures. . 1971. A new species of Clypeaster (Echinodermata, Echinoidea) from San Felix Island, with a key to the recent species of the Eastern Pacific Ocean.—Pacific Science 25(2): 165-170, 4 figures. (DLP) Department of Invertebrate Zoology, National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560; (TFP) Oregon Institute of Marine Biology, Charleston, Oregon 97420. PROC. BIOL. SOC. WASH. 92(4), 1979, pp. 801-803 HYMENASTER KIERI, A NEW SPECIES OF STARFISH OF THE FAMILY PTERASTERIDAE (ECHINODERMATA: ASTEROIDEA) Maureen E. Downey Abstract.—A new starfish, Hymenaster kieri, is described from 2,625 m off Virginia, bringing to 12 the known Atlantic species of Hymenaster. During a cruise by the Virginia Institute of Marine Science on R/V Gillis in November 1974, a large and handsome specimen of the starfish genus Hymenaster was collected in deep water off the coast of Virginia by Kathy Larson and Michael Sweeney, of the National Museum of Natural History. Specimens of Hymenaster are apparently widely distributed in the world’s oceans, at lower bathyal and abyssal depths, but are relatively rare in col- lections because of the difficulties of collecting such soft-bodied animals from great depths. Family Pterasteridae Perrier, 1875 Genus Hymenaster Thompson, 1873 Hymenaster kieri, new species Type.—USNM E 18204 (Holotype), 36°44’N, 73°45'W, 2,625 m. Etymology.—This species is named in honor of Dr. Porter M. Kier on the occasion of his resignation as Director, National Museum of Natural His- tory. Diagnosis.—Form broadly stellate, inflated, fleshy; arms 5, broad-based, petaloid, with tips tapering rapidly to subacute point; abactinal surface with 9 regular rows of paxillae on each arm; paxillae rather small, with 3—4 basal lobes slightly flattened and truncate; pedicel short, bearing | very large, spike-like spine elevating a tent of dorsal membrane; interradially, a broad, bare, fleshy marginal flange; abactinal spiracles tiny, numerous, in irregular, meandering groups; centrally, 5 large valves of 10 equal, blunt spines webbed together surrounding large osculum; prominent circle of spines in groups of 3—5 outside valves; within the osculum an enormous bulbous madreporite is visible as well as a small pavement of stout, irregular, flat plates; interbrachial septae a sheet of tissue in which are embedded large flat plates which, from their shape, seem obviously derived from paxillae; actinally, ca. 45 actinolateral spines, well-spaced, not very long, broad- based, with blunt tips, the first 20-30 of nearly equal length, becoming rapidly shorter, more crowded beyond interradial marginal flange, at arm 802 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 1. Hymenaster kieri. tip; these spines, embedded in actinal membrane, do not meet interradially; ambulacral grooves wide, petaloid; adambulacral plates elongate, with distal half underlying next adambulacral plate, proximal half bearing 2-3 short, stout, conical spines in oblique row, and a rounded, ridged opercular spine covering the segmental apertures; mouth plates plowshare-shaped, bearing 1 large, conical oral spine, a similar suboral spine, and 2-3 small, finer lateral spines. R = 130 mm, r = 85 mm, R = 1.5. Color: Livid pink abactinally, blood red actinally. VOLUME 92, NUMBER 4 803 Discussion.—Because of its thick, fleshy tegument, this species seems to be closely allied to Hymenaster carnosus Sladen (1882), from the South Atlantic. It differs in having 9 regular rows of paxillae bearing one spine each (except those encircling the osculum), vs. an irregular arrangement of paxillae with an unknown number of spines; petaloid ambulacral grooves, vs. grooves of uniform width; 2-3 parallel conical adambulacral spines, vs. 2 oblique needle-like spines; adambulacral spines bare, vs. ensacculated; operculate spine covering segmental aperture, vs. fleshy pad; 2 lateral mouth spines, vs. 3 or 4; actinolateral spines approximately 45 and well-spaced, vs. 50-60 closely placed spines; spiracles very numerous, in meandering, ill-defined groups, vs. 2 or more in small round groups. A tabular key to the Atlantic species of Hymenaster was published by Sibuet in 1976, and 2 new species were also added to the genus. The only comprehensive reviews of the genus were by Sladen in 1882 and 1889, in reports on the Challenger asteroids. Prior to the Challenger expedition, only 4 species of Hymenaster were known from the Atlantic; the Challenger collections added 3 new Atlantic species. With the addition of Hymenaster kieri, the number of Hymenaster species now known from the Atlantic is 12. Literature Cited Sibuet, Myriam. 1976. Le genre Hymenaster (Asterides) dans |’océan Atlantique.—Bull. Mus. Nat. Hist. Natur., ser. 3, no. 368, Zool. 256:305—324, figs. 1-7. Sladen, W. Percy. 1882. The Asteroidea of H.M.S. Challenger Expedition (preliminary no- tice).—Jour. Linn. Soc. (Zool.), 16: 189-246. . 1889. Report of the Asteroidea collected by H.M.S. Challenger.—Repts. Sci. Res. H.M.S. Challenger, Zoology 30:893, 117 pls. Department of Invertebrate Zoology, National Museum of Natural His- tory, Smithsonian Institution, Washington, D.C. 20560. PROC. BIOL. SOC. WASH. 92(4), 1979, pp. 804-811 A NEW CRAYFISH FROM THE OUACHITA RIVER BASIN IN ARKANSAS (DECAPODA: CAMBARIDAE) Horton H. Hobbs, Jr. Abstract.—Procambarus (Girardiella) reimeri is described and illustrated from specimens collected in the Ouachita River basin in Polk County, Ar- kansas. Its affinities to P. (G.) gracilis, P. (G.) liberorum, and to members of the subgenus Austrocambarus are discussed. During April 1973, Jean E. Pugh, Georgia B. Hobbs, and I spent several days in western Arkansas collecting crayfishes. As a result of heavy rains the streams were at flood stage; therefore most of our efforts were expended in collecting inhabitants of roadside ditches and burrows. Two previously undescribed members of the genus Fallicambarus obtained at that time were described in the same year (Hobbs, 1973). Attention to new representatives of the subgenus Girardiella (genus Procambarus), however, was deferred, for it was my intention to offer these specimens to Rollin D. Reimer who had concentrated his attention on this crayfish group. Inasmuch as some years have elapsed since his unpublished dissertation was completed and his interests have broadened to include other crayfish groups, I offer the description of this new species, the existence of which appears not to have been known to Dr. Reimer. I take pleasure in naming it in his honor. I extend my thanks to Dr. Pugh and to my wife for their assistance in collecting the material on which this description is based and to Fenner A. Chace, Jr., Margaret A. Daniel, and C. W. Hart, Jr., colleagues at the Smithsonian Institution, for their criticisms of the manuscript. Procambarus (Girardiella) reimeri, new species Fig. 1 Diagnosis.—Body pigmented, eyes well developed. Rostrum without marginal spines and median carina. Carapace lacking cervical spine or tu- Fig. 1. Procambarus (G.) reimeri (a-c, e-g, i-k, m-p, and r from holotype; d, h fon morphotype, and /, g from allotype): a, Lateral view of carapace; b, Dorsal view of telson and uropods; c, d, e, Mesial view of first pleopod; f, Caudal view of first pleopods; g, h, i, Lateral view of first pleopod; j, Epistome; k, Cephalic view of distal part of first pleopod; /, Annulus ventralis; m, Caudal view of distal part of first pleopod; n, Antennal scale; 0, Dorsal view of carapace; p, Basal podomeres of third, fourth, and fifth pereiopods; g, r, Dorsal view of distal podomeres of cheliped. 805 VOLUME 92, NUMBER 4 806 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON bercle. Areola 16.3 to 25.3 (average 20.5) times as long as broad and con- stituting 34.9 to 40.2 (average 37.5) percent of total length of carapace (41.3 to 44.9, average 43.9, percent of postorbital carapace length). Suborbital angle weak but distinct. Postorbital ridge lacking cephalic spine or tubercle. Hepatic area tuberculate; branchiostegal spine vestigial or absent. Antennal scale about, or slightly more than, twice as long as wide. Ischium of third pereiopod of first form male with simple hook overreaching basioischial articulation; hook not opposed by tubercle on corresponding basis; coxae of fourth and fifth pereiopods lacking caudomesial boss. First pleopods of first form male reaching coxae of third pereiopods, symmetrical, bearing proximomesial spur and cephalic shoulder at base of terminal elements, lacking subterminal setae but with cephalomesial row of setae along distal third to half of appendage; terminal elements (all at least partly cornified) consisting of (1) long, tapering, slightly curved mesial process extending distally beyond other terminals; (2) small, acute, slightly curved, but distally directed cephalic process at cephalic base of mesial process; (3) rather in- conspicuous but composite central projection arising from between bases of cephalic, mesial, and caudal processes, and closely associated with latter; and (4) very prominent caudal process subacute and obscuring central pro- jection in lateral aspect and somewhat rounded distally and flattened cau- dally in caudal aspect. Female with annulus ventralis about 1.5 times as broad as long, subsymmetrical in outline, with cephalomedian trough lead- ing to sigmoid sinus, latter ending near median line anterior to caudal margin of annulus; ridges flanking trough devoid of tubercles; preannular (= proan- nular, Hobbs, 1967:9) plate well developed; postannular sclerite almost 0.9 times as long and wide as annulus, broadly elevated ventrally; first pleopod present. Holotypic male, form I.—Cephalothorax (Figure la, 0) subovate, weakly compressed laterally, although greatest width of carapace slightly more than height at caudodorsal margin of cervical groove (16.3 and 15.7 mm). Ab- domen distinctly narrower than thorax (12.2 and 16.3 mm). Areola 17.9 times as long as broad; punctations sparse, only | or 2 across narrowest part. Cephalic section of carapace almost 1.7 times as long as areola, latter comprising 37.7 percent of total length of carapace (44.2 percent of post- orbital carapace length). Surface of carapace punctate dorsally, mostly gran- ulate laterally, granules replaced by tubercles in hepatic region. Rostrum broad basally with weakly convergent margins turning rather suddenly an- teromesially over penultimate podomere of antennule and ending in slightly upturned tip above distomesial end of podomere; margins not conspicuously thickened; upper surface concave with rows of submarginal punctations, very few others in anterior third, but more abundant and larger posteriorly. Subrostral ridges weak and evident only along caudodorsal margin of orbit. Postorbital ridges moderately prominent, low but swollen caudally and VOLUME 92, NUMBER 4 807 rounded cephalically, lacking tubercles or spines. Suborbital angle weak but well defined. Branchiostegal and cervical spines absent. Abdomen shorter than carapace (30.0 and 33.2 mm). Pleura of third through fifth segments subtruncate ventrally and rounded caudoventrally. Cephalic section of telson with 2 spines in each caudolateral corner, lateral ones fixed (Fig. 1b). Cephalic lobe of epistome (Fig. 1/) broadly triangular and with irregularly arranged groups of plumose setae marginally; margins not markedly thickened and ventral surface subplane; main body of epi- stome lacking median fovea, only slightly depressed; epistomal zygoma broadly arched. Ventral surface of proximal podomere of antennular pe- duncle with spine short distance distal to midlength. Antennal peduncle lacking spines on basis and ischium; flagellum reaching tergum of second abdominal segment. Antennal scale (Fig. In) twice as long as broad, widest distal to midlength; greatest width of lamellar area about 2 times that of thickened lateral part. Third maxilliped extending to level of base of ultimate podomere of an- tennule, ventral surface largely hidden by tufts of long plumose setae, those concealing lateral half of ventral surface of ischtum borne in submarginal row and single distal cluster; part of those on mesial half long and plumose and others stiff and simple. Merus completely obscured in lateral aspect by long plumose setae extending distally from ischium and from ventrolateral surface of merus. Right chela (Fig. Ir) subovate in cross section, not strongly depressed; palm approximately 1.4 times as broad as length of mesial margin; latter slightly greater than one-third total length of chela; almost entire palm stud- ded with squamous to subsquamous tubercles, those situated proximolater- ally smaller and more withdrawn than more mesial and distal ones. Mesial surface of palm with row of 6 tubercles flanked dorsally and ventrally by irregularly arranged ones; ventral surface with | tubercle on ridge flanking base of dactyl. Both fingers with low longitudinal median ridges dorsally and ventrally; ridges flanked proximally by squamous tubercles and distally by punctations. Opposable margin of fixed finger with row of 7 tubercles (third from base largest) along proximal half and very large tubercle pro- jecting from lower level in distal part of penultimate fourth; single row of minute denticles, interrupted by tubercular row, extending from proximal- most tubercle to corneous tip of finger. Opposable margin of dactyl deeply excised proximally, with row of 5 tubercles (third from base conspicuously larger than others) along proximal two-thirds, and minute denticles arranged as on fixed finger; mesial margin with row of 8 tubercles (decreasing in size distally) along proximal two-thirds of finger. Carpus of cheliped longer than broad with oblique furrow dorsally; mesial surface with | very large conical tubercle and 3 proximal to it; few tubercles on dorsomesial surface, otherwise punctate except for tubercles on disto- 808 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON ventral margin: 1 on mesiodistal condyle, and row of 5 (decreasing in size laterally) lateral to that on condyle. Merus tuberculate along dorsodistal two-thirds of podomere; ventral sur- face with mesial row of 15 (left with 16) tubercles and 10 in lateral; otherwise podomere mostly punctate. Ischium with row of 4 (left with 3) tubercles ventromesially. Hook on ischium of third pereiopod (Fig. 1p) simple, overreaching bas- ioischial articulation, not opposed by tubercle on corresponding basis. Cox- ae of neither fourth nor fifth pereiopods with caudomesial boss. Sternum between third, fourth, and fifth pereiopods rather shallow but ventrolateral margins bearing conspicuous fringe of plumose setae obscuring first pleopods. First pleopods (Fig. Ic, e-g, i, k, m) as described in ‘Diagnosis.’ In addition, proximomedian lobe very long. Uropods (Fig. 1b) with both lobes of basal podomere bearing short acute spines; mesial ramus with distomedian spine small and situated proximal to distal margin. Allotypic female.—Differing from holotype, other than in secondary sex- ual features, in following respects: cephalic lobe of epistome devoid of plu- mose setae, fovea on main body distinct; submarginal setae on ischium and merus of third maxilliped much reduced so that much of surface of lateral half of ischium exposed; opposable margin of fixed finger of chela (Fig. 1q) with row of 8 tubercles, that of dactyl also with 8, mesial margin of latter with row of 10; ventromesial row of tubercles on merus of cheliped con- sisting of 18 (17 on left) and ventrolateral row of 12; 3 tubercles forming row on basis of same appendage; sternum moderately deep but ventrolateral margins lacking conspicuous setae. (See Tab. 1.) Annulus ventralis (Fig. 1/) as described in ‘*Diagnosis.”’ Morphotypic male, form II.—Differing from holotype in following re- spects: branchiostegal spine vestigial; cephalic lobe of epistome more broad- ly triangular and with fewer marginal setae; lateral row of submarginal setae on ischium of third maxilliped reduced, but appendage more hirsute than that of allotype; opposable margin of fixed finger of chela with row of 8 tubercles, that of dactyl with 6; ventromesial margin of merus of cheliped with row of 12 (13 on left), and ventrolateral row of 13 on right and 10 on left; ischium of third pereiopod not nearly so well developed and not reach- ing basioischial articulation. (See Tab. 1.) First pleopod (Fig. ld, h) with mesial process proportionately more robust than that of holotype; cephalic process reduced to short acute tubercle; central projection and caudal pro- cess not so large but more clearly defined; setae on distal half of appendage much less conspicuous. Color Notes (Based on topoparatypic male, form I).—Carapace pinkish cream, dorsolateral part of branchiostegites suffused with brown; posterior VOLUME 92, NUMBER 4 809 Table 1.—Measurements (mm) of Procambarus (G.) reimeri. Holotype Allotype Morphotype Carapace: Entire length 53.2 35,0 29.9 Postorbital length 28.3 31.0 24.9 Width 16.3 Wed 13.9 Height 15-7 17.2 sy Areola: Width 0.7 0.8 0.5 Length j WAS) 13.9 10.8 Rostrum: Width ye 5.6 5.0 Length 6.3 6.1 Sey Chela: Length, palm mesial margin 8.2 S27 5.6 Palm width 11.8 8.5 8.0 Length, lateral margin 27:1 20.5 18.0 Dactyl length L7:2 13.4 11.6 Abdomen: Width 1222 14.4 10.4 Length 30.0 3313 26.0 part of mandibular adductor region, areola, and posterodorsal region pinkish brown. First abdominal tergum dark brick red, second through fifth terga cherry red, and sixth, together with all pleura, telson, and uropods, similarly colored but suffused with tan. Antennule olive mottled with pinkish tan; antennal peduncles paler and more pinkish than olive; flagella of both ap- pendages olive tan. Cheliped pinkish cream from base to proximal half or two-thirds of merus, where becoming olive tan to brown; latter extending Over carpus; chela olive mesially and dorsally, and pinkish orange ventrally and laterally, colors merging dorsolaterally. Remaining pereiopods pinkish cream proximally becoming diffused with olive and distinctly pale olive with reddish brown setae distal to midlength of merus. Type-locality.—Burrows in roadside ditch about five miles northeast of Mena, Polk County, Arkansas, on unnumbered road to Irons Fork River. The comparatively simple burrows, one-half to one meter in depth, were constructed in a sandy clay soil. Disposition of types.—The holotype, allotype, and morphotype are de- posited in the National Museum of Natural History (Smithsonian Institu- tion), nos. 148880, 148881, and 148882, respectively, as are the paratypes consisting of 3 6 I,4 6 II, 10 2,51j 5, 48j @. Size.—The largest specimen available is a female having a carapace length 810 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON of 41.5 (postorbital carapace length 36.0) mm. The largest and smallest first form males have corresponding lengths of 39.4 (33.9) mm and 31.6 (26.6) mm. Females carrying eggs or young have not been collected. Range and specimens examined.—All of the known localities are in the Ouachita River basin of Polk County, Arkansas. The limited series available was collected by Georgia B. Hobbs, Jean E. Pugh, and H.H.H., Jr. Type- locality, 2 6 I, 1 6 II, 2 2, 1j 6, 18 Apr 1973; roadside ditch about 7 mi NE of Mena on unnumbered road to Irons Fork River, 4j 6, 1j 2, 18 Apr 1973; roadside ditch adjacent to Ouachita River, 3.4 mi E of junction of US Hwy 71 on St Rte 88, 2 6 1,2 6 Hl, 2 2, 20j 3, 23j 2; roadside pool on Posey Hollow Road 4.5 mi N of Acorn and 1.4 mi E of US Hwy 71, 2 j 6, 20 Apr 1973; flooded tributary to Irons Fork on Posey Hollow Road about 1 mi E of US Hwy 71, 1j 2, 20 Apr 1973; pool in roadside ditch 3.1 mi SE of St Rte 5 on St Rte 375, 2 6 Il, 7 2, 243 5, 233 2, 20 Apr 1973. Variations.—The variations noted are rather insignificant except for an occasional ratio such as that between areola length and carapace length; it is as low as 34.9 percent in one of the females, but the closest approached by another specimen was 36.0 percent, and the average is 37.5. The ce- phalomedian lobe of the epistome may or may not bear a marginal row of plumose setae. The antennae reach caudally to the second or third abdom- inal tergum. Pubescence on the third maxilliped is highly variable but seems to be longer and more conspicuous in the first form males; in some individ- uals of both sexes, the submarginal row may be short, obscuring little of the ventral surface of the ischium. The tubercular row on the mesial surface of the palm consists of five to eight, that on the opposable margin of the fixed finger seven to 11, that on the dactyl five to seven, that on the ven- trodistal margin of the carpus two to six, that in the ventrolateral row on the merus 10 to 13, and in the ventromesial row 13 to 16. In the juvenile females the preannular plate is not evident, and the postannular sclerite is proportionately smaller than in the adult females. Relationships.—Procambarus (G.) reimeri has its closest affinities with P. (G.) gracilis (Bundy, 1876:5) and P. (G.) liberorum Fitzpatrick (1978:533). It differs from the former most conspicuously in possessing a broader areola, in the shorter central projection (not overreaching the caudal process) of the first pleopod of the male, and in the absence of tubercles on the cephalolateral ridges of the annulus ventralis. It may be distinguished from P. (G.) liberorum also in having a broader areola, in the disposition of the caudal process (appearing acute in lateral aspect), and in lacking tubercles on the annulus ventralis. In all of the adult females of P. (G.) reimeri, the annulus ventralis is separated from the V-shaped sternum an- terior to it by a firm though comparatively weakly calcified, ventrally convex structure that I interpret as being a homologue of the preannular plate char- acteristic of members of the subgenus Austrocambarus. To be sure, com- VOLUME 92, NUMBER 4 811 paratively the plate in P. (G.) reimeri is not nearly so large as that in the crayfishes of Cuba, southern Mexico, Guatemala, and Honduras, but the similarities in position and texture are suggestive of a feature shared in common. If these may be assumed to be homologous structures, further credence can be given to previously postulated (Hobbs, !967:11) close af- finities between the members of Austrocambarus (derived from the Mexi- canus-Cubensis ancestral stock) and those of Girardiella (Graciloid stock). Literature Cited Bundy, W. F. 1876. In S. A. Forbes, List of Illinois Crustacea, with descriptions of new species.—Bull. Ill. Mus. Nat. Hist. 1:3, 4, 5, 24-25. Fitzpatrick, J. F., Jr. 1978. A new crawfish of the subgenus Girardiella, genus Procambarus from northwest Arkansas (Decapoda, Cambaridae).—Proc. Biol. Soc. Wash. 91(2):533- 538, | fig. Hobbs, Horton H., Jr. 1967. A new crayfish from Alabama caves with notes on the origin of the genera Orconectes and Cambarus (Decapoda: Astacidae).—Proc. U. S. Nat. Mus. 123(3621):1-17, 21 figs. . 1973. New species and relationships of the members of the genus Fallicambarus.— Proc. Biol. Soc. Wash. 86(40):461-481, 4 figs. Reimer, Rollin Dewayne. 1969. A taxonomic study of the Gracilis Section of the Genus Procambarus.—Unpublished dissertation, Tulane University, New Orleans, Louisiana. 190 pages. Department of Invertebrate Zoology, Smithsonian Institution, Washing- ton, D. C. 20560. PROC. BIOL. SOC. WASH. 92(4), 1979, pp. 812-813 THE PROSTOMIAL PIT IN BOTHRIONEURUM VEJDOVSKYANUM STOLC (OLIGOCHAETA): A NOTE ON DETAIL REVEALED BY SEM Peter M. Chapman Abstract.—Two SEM views of the prostomial pit in Bothrioneurum vej- dovskyanum Stolc show that it contains cilia. The genus Bothrioneurum of the Tubificidae (Oligochaeta) is partly de- fined by the presence of a dorsal ciliated pit in its prostomium (Brinkhurst and Jamieson, 1971). The purpose of this study was to illustrate the structure of this diagnostic feature by photographing the type-species, B. vejdov- skyanum Stolc, under a scanning electron microscope. Worms preserved in formalin were dehydrated with alcohol in 10% steps and brought up to 100% amyl acetate. The worms were then critical point dried, gold coated and examined under a JEOL JSM-35 scanning electron microscope. Two views of the prostomial pit are presented. Figure | shows the position of the pit in the prostomium. Figure 2 is a high magnification (2,000) view of the pit showing it to contain cilia. The prostomial pit is therefore shown to be a clearly defined structure containing cilia, which suggests that the pit has a chemosensory function. Literature Cited Brinkhurst, R. O. and B. G. M. Jamieson. 1971. Aquatic Oligochaeta of the world.—Univer- sity of Toronto Press, Toronto, 860 pp. Biology Department, University of Victoria, PO Box 1700, Victoria, B.C.., Canada, V8W 2Y2. 813 VOLUME 92, NUMBER 4 Anterior region of Bothrioneurum vejdovskyanum. Note the pit in the center of the Fig. 1. prostomium. 150x. Fig. 2. Prostomial pit of Bothrioneurum vejdovskyanum. 2,000x. PROC. BIOL. SOC. WASH. 92(4), 1979, pp. 814-836 NEW SPECIES OF ANTHURIDEANS FROM THE COOK AND FIJI ISLANDS (CRUSTACEA: ISOPODA: ANTHURIDEA) Brian Kensley Abstract.—Seven new anthuridean isopod species are reported from the Fiji and Cook islands. These include five anthurids, viz. Apanthura mana, Mesanthura childi, Natalanthura fijiensis, Panathura collaris, Pendanthura rarotonga, and two paranthurids, Paranthura astrolabium and Paranthura melanesica. During July-August 1978, Mr. C. A. Child of the Smithsonian Institution spent ten days doing shallow-water collecting in the Cook and Fiji islands. Various habitats, including coral and calcareous algal rubble, and algal mats were sampled and a total of 34 anthuridean specimens collected. These anthurideans proved to belong to seven species, all of which are new. This is hardly surprising, as the only reports of anthurids from the central Pacific are those of Miller and Menzies (1952) from Hawaii (whose report deals with four new species) and Stebbing (1900) who described two new species from the Loyalty Islands. These reports of new species indicate how poorly the central Pacific has been collected, and also support the idea that although genera are often widespread, anthuridean species in shallow water habitats tend to have restricted geographical ranges. Family Anthuridae Apanthura mana, new species Figs. 1-2 Description.—¢. Integument not indurate, lacking pigmentation. Body proportions: C << 1>2>3<4<5>6>7. Cephalon with low rostral point barely extending beyond rounded anterolateral corners; eyes small, dorsolateral. Pereonites 4, 5, and 6 each with shallow middorsal pit. Pleo- nites 1-5 fused; 6 free, with posterodorsal margin convex. Telson dorsally evenly convex, proximal half parallel-sided, distal half tapering to bluntly rounded setose apex, with narrow hyaline margin; 2 large basal statocysts present. Antennular peduncle 4-segmented, basal segment subequal to 3 distal seg- ments together; flagellum of 2 articles. Antennal peduncle 5-segmented, sec- ond segment strongly grooved to accommodate antennule; flagellum of 3 articles, 2 distal articles tiny. Mandibular palp 3-segmented, distal segment VOLUME 92, NUMBER 4 815 Fig. 1. Apanthura mana: a, Female in dorsal view; b, Maxilla; c, Mandible; d, Maxilliped: e, Pereopod 1; f, Antennule; g, Antenna. 816 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON ZZ Ze Ze KE CLA KK > Fig. 2. Apanthura mana: a, Telson and uropod; b, Pereopod 2; c, Pereopod 7; d, Pleopod 1. with 2 terminal setae; segment 2 almost 3 times length of segment 3; incisor of 3 cusps; lacinia with 4 serrations; molar bluntly rounded. Maxilla with 6 distal spines. Maxilliped 5-segmented, terminal segment set obliquely at outer distal corner of segment 4, with 5 simple setae; thin-walled endite on inner surface almost reaching end of segment 3. Pereopod | subchelate, VOLUME 92, NUMBER 4 817 unguis slightly more than half length of rest of dactylus; propodus expanded, palm with slight notch at midpoint and few simple setae; carpus triangular, distally rounded. Pereopod 2 unguis about one-third length of rest of dac- tylus; propodus with strong serrate sensory spine at posterodistal angle. Pereopods 4—7, posterior margins of propodi and dactyli with fringed scales; propodi serrate sensory spine at posterodistal angle; carpi triangular, with short anterior margin, short sensory spine at posterodistal angle. Pleopod 1 exopod operculiform, with numerous distal plumose setae, endopod some- what shorter than and one-third width of exopod, with 5 distal plumose setae; basis with 3 retinaculae. Uropodal exopod elongate/oval, just reach- ing endopod, with slight subapical notch in outer margin, numerous marginal plumose setae and few simple setae; endopod reaching beyond telsonic apex, tapering to broadly rounded setose apex. Etymology.—The species was collected off Mana Island in the Fiji group, hence the specific name. Material.—Holotype, USNM 172271, 2°, 3.9 mm. Paratypes, USNM 172272, 2 2, 3.2 mm, 3.9 mm. Mana Island, Fiji; taken from coral rubble in 1 m water on fringing reef. Remarks.—The genus Apanthura, containing as it does many very similar species, undoubtedly requires revision. Differences between species of Apanthura are usually subtle, and in the present case, several species agree with A. mana in numerous features. A. sandalensis Stebbing from the Loy- alty Islands is a larger species (7 mm) with a strongly notched uropodal exopod. A. inornata Miller and Menzies from Hawaii is of similar size (3.75 mm) but has a more strongly notched propodal palm of pereopod 1, and the uropodal exopod is relatively broader. Miller and Menzies (1952:4, fig. If) mention and figure a ‘‘single toothed bristle in the setal row’’ of the mandible of A. inornata. This is an extremely unusual feature and would make the Hawaiian species unique in this regard. Mesanthura childi, new species Figs. 3-4 Description.—. Integument not indurate. Body proportions: C = | > 2<3<4=5>6>7. Cephalon with low rostrum not extending beyond anterolateral corners; large middorsal pigment patch; small dorsolateral eyes present. Pigment pattern: 2 proximal patches on pereonites | and 2, 3 on pereonite 3, 2 distal patches on pereonites 4-7, 2 patches overlapping ple- onites 5 and 6. Telson widest at about midpoint, apex broadly rounded with cluster of simple setae; 2 large proximal statocysts present. Antennular peduncle 4-segmented, basal segment slightly shorter than 3 distal segments together; flagellum of 2 articles, terminal article with 3 aesth- etascs. Antennal peduncle 5-segmented, segment 2 grooved to accommo- 818 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Ki Fig. 3. Maxilla; e, Maxilliped; f, Telson and uropod. Mesanthura childi: a, Female in dorsal view; b, Cephalon of male; c, Mandible; d, date antennule; flagellum of 2 articles. Mandibular palp 3-segmented, seg- ment 2 twice length of terminal segment; latter with 3 distal spines; incisor with 3 rounded cusps; lacinia with 4 teeth; molar bluntly rounded. Maxilla with 1 strong and 6 smaller distal spines. Maxilliped 5-segmented, terminal VOLUME 92, NUMBER 4 819 Tl PRD Ke SS z UR SQ 4 Fig. 4. Mesanthura childi: a, Pleopod 1; b, Pleopod 2 ¢; c, Pereopod | 2; d, Pereopod | 36; e, Pereopod 2; f, Pereopod 7. segment with 5 setae, set at outer distal corner of segment 4; endite on inner surface reaching to middle of segment 3, with single terminal seta. Pereopod 1 subchelate; unguis one-third length of dactylus, with short accessory spine 820 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON at base; propodal palm slightly concave, with few simple setae; carpus tri- angular, distally rounded, with few simple setae. Pereopod 2 propodus bare- ly expanded proximally, with strong serrate sensory spine at posterodistal corner. Pereopods 4—7 unguis about one-third length of dactylus, latter with few fringed scales on posterior margin; propodus with 2 serrate sensory spines at posterodistal corner, several fringed scales on posterior margin; carpus triangular, anterior margin shorter than posterior margin. Pleopod 1 exopod operculiform, with plumose setae on distal margin; endopod about one-third width of and almost as long as exopod, with 4 distal plumose setae; basis with 3 retinaculae. Uropodal exopod oval, with shallow notch in distal margin, reaching beyond basis, with numerous marginal plumose setae; endopod distally rounded, with numerous simple setae; reaching be- yond telsonic apex. 6. Cephalon with eyes much larger than °. Antennular flagellum elon- gate, of 6 articles bearing filiform aesthetascs. Pigment pattern similar to ? but more diffuse, extending onto ventral surface. Pereopod 1 unguis half length of dactylus; propodal palm straight, with numerous setae on inner surface. Pleopod 2 copulatory stylet on endopod extending beyond both rami, distally rounded. 7 Etymology.—The species is named for Mr. C. A. Child of the Department of Invertebrate Zoology, Smithsonian Institution, who collected the entire anthurid collection described in this paper. Material.—Holotype, USNM 172273, 2, 3.0 mm; Allotype, USNM 172274, 6, 2.6 mm. Mana Island, Fiji; taken from coral rubble in 1 m water on fringing reef. Remarks.—Mesanthura childi is one of the smallest species of this genus yet recorded. The development of the copulatory stylet of the male indicates that these are mature specimens. Only two species of Mesanthura have been recorded from the Pacific, viz. M. occidentalis Menzies and Barnard, from southern California, and M. hieroglyphica Miller and Menzies, from Hawaii. Neither of these possess a pigment pattern at all similar to the present species. Since pigment pattern is regarded as fairly constant and species-distinct in Mesanthura the present material must be seen as hitherto undescribed. Natalanthura fijiensis, new species Figs. 5-6 Description.—¢. Integument moderately indurate, strongly pitted. An- terolateral corners of cephalon extending slightly beyond low rostrum, eyes well developed, pigmented. Pereonites dorsally and laterally pitted. Body proportions: C<1<2>3<4=5>6>7. Pleonites 1-S fused, with transverse rows of pits marking lines of fusion; pleonite 6 indistinguishably VOLUME 92, NUMBER 4 821 Fig. 5. Natalanthura fijiensis: a, Holotype in dorsal view; b, Mandible; c, Maxilliped; d, Maxilla; e, Antennule; f, Antenna; g, Uropod. fused with telson. Telson distally evenly rounded, with strong longitudinal mediodorsal ridge flanked by pits; margin entire; statocysts difficult to de- tect, probably present. Antennular peduncle 4-segmented; flagellum of 2 articles. Antennal pe- duncle 5-segmented, segment 2 grooved to accommodate antennule; flagel- lum of 3 articles. Mandibular palp 3-segmented, middle segment about twice length of distal segment; latter with 3 distal fringed spines; incisor of 2 blunt cusps; lacinia narrow, with 11 serrations; molar spike-like, elongate, finely ridged. Maxilla slender, with 4 distal spines. Maxilliped 5-segmented, seg- 822 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON \ 7} { )) by \ + 4) K ; HAVA PANY ix y Wali / f i) . | Y \ NIN \\ 7h ( u /, \ \’ i 7 y ‘) { VANS / y py 4 \ / SS EE LEE a Fig. 6. Natalanthura fijiensis: a, Pleopod 1; b, Pereopod 7; c, Pereopod 1. ment 2 relatively elongate, terminal segment semicircular, with 4 distal sim- ple setae; endite on internal surface extending to midlength of segment 4. Pereopod | not differing markedly from following pereopods; only slightly more robust; unguis about half length of dactylus, with strong spine at base; propodus with strong posterodistal sensory spine; carpus short, triangular. Pereopods 4—7 propodus slightly more slender than in anterior pereopods, with strong posterodistal sensory spine and 2 elongate fringed spines dis- tally; carpus roughly triangular, with short anterior margin, sensory spine posterodistally. Pleopod 1 basis with 3 retinaculae; endopod subequal in length and about half width and slightly overlapped by exopod; both rami with several elongate plumose setae, together forming operculum. Uropodal exopod elongate/oval, with simple marginal setae; basis shorter than en- dopod; latter elongate/oval, with relatively elongate marginal simple setae; both rami with broad hyaline entire margin. VOLUME 92, NUMBER 4 823 Etymology.—The species is named for the Fiji Islands group, whence it was collected. Material.—Holotype, USNM 172275, 2, 3.4 mm. South-west coast of Viti Levu, Fiji; taken from coral rubble from lagoonal shallows behind bar- rier reef, in 1 m of water. Remarks.—The present species may easily be separated from the two previously described species by the pattern of dorsal integumental pits, but also by several other characters tabulated below: Table N. natalensis N. sp. (Belize) N. fijiensis (Kensley, 1978) (Kensley, in press, a) Pereopod 7 Present Present Absent Antennular flagellum articles 2 3 3 Antennal flagellum articles 3 6 f Lacinia serrations 1] 12 7 Telsonic margin Entire Serrate Entire Both the present species and Natalanthura sp. from Belize were taken from shallow-water coarse coral reef sediments consisting mainly of coral rubble, while the Natal species was taken from much deeper water, but also from coarse sediments of animal origin. Panathura collaris, new species Figs. 7-9 Description.—°?. Integument thin, not indurate. Body proportions: C < 1>2<3=4=2=5>6>7. Cephalon with U-shaped band of red-brown pigment dorsally; eyes dorsolateral. Pereonite 7 with posterolateral lobes overlapping anterior pleonites laterally. Pleonites free, 1—4 short, subequal; pleonite 5 twice length of pleonite 4, with fringe of dorsolateral plumose setae; pleonite 6 twice length of 5, with rounded middorsal lobe on posterior margin. Telson distally broadly and evenly rounded, with broad hyaline margin, dorsal surface faintly concave, with scattered short setules, ventral surface convex; distal margin with several elongate simple setae. Antennular peduncle 4-segmented, basal segment broad, as long as remain- ing segments together; flagellum of single article, bearing elongate simple setae. Antennal peduncle 5-segmented, segment 2 longest and broadest; flagellum of 2 short setiferous articles. Mandibular palp 3-segmented, middle segment about twice length of segments | and 3; latter with 4 distal fringed spines; incisor of 3 cusps; lacinia with 4 large serrations and several fine proximal spinules; molar lacking. Maxilla slender, tapering distally with | strong and 4 smaller spines. Maxilliped 6-segmented, with large endite reaching to segment 5; segment 2 short; 4 simple setae and short fringed 824 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 7. Panathura collaris: a, Female in dorsal view; b, Mandible; c, Maxilla; d, Maxilliped; e, Antennule °; f, Antenna; g, Antennule ¢; h, Pereopod 7. scale distally on terminal segment. Pereopod 1 unguis almost one-quarter length of entire dactylus; propodus broad, palm straight, with thin trans- parent margin produced into triangular process proximally; strong fringed spine on inner surface near dactylar articulation; carpus triangular, with thin VOLUME 92, NUMBER 4 825 NY EEX ~~ Ys ~! \N x SOO LV) \ é x 2>3>46>/7. Cephalon with prominent rostrum extending beyond anterolateral lobes. Eyes dorsolateral. Brood pouch formed by 3 pairs of oostegites on pereo- nites 3-5. Pleonites 1-5 fused; 6 free, with fringe of plumose setae. Telson basally broad, with 2 large statocysts in proximal half, distally evenly round- ed. Antennular peduncle 3-segmented, basal segment longest; flagellum of 3 short articles. Antennal peduncle 5-segmented; flagellum of 1 short article. Mandibular palp reduced to small papilla bearing 1 terminal and | subter- minal simple seta; incisor of 3 rounded cusps; lacinia with 8 marginal ser- rations; molar process triangular. Maxilla slender, with single strong ter- minal spine and 7 small spines. Maxilliped 3-segmented, terminal segment with 6 simple setae on distal margin; thin-walled endite tipped with single seta. Pereopod | unguis more than half length of dactylus, with strong spine at base; propodus proximally broad, palm sinuous, with few simple setae; group of 8 fringed spines on inner distal surface; carpus short, triangular. Pereopod 2 far less robust than pereopod 1, propodus with strong postero- distal fringed sensory spine. Posterior pereopods, propodus with fringed scales on posterior margin, strong sensory spine posterodistally; carpus tri- angular, with short anterior margin. Pleopod | exopod operculiform, as long as endopod, both with plumose setae marginally, reaching to distal margin of basis; endopod oval, with simple and plumose setae. 828 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 10. Pendanthura rarotonga: a, Female in dorsal view; b, Mandible; c, Maxilla; d, Maxilliped; e, Antennule; f, Antenna; g, Telson and uropod. Etymology.—The species is named for Rarotonga Island in the Cook Is- land group, where it was collected. Material.—Holotype, USNM 172281, ovigerous 2, 3.2 mm; Paratypes, USNM 172282, ovigerous °, 3.0 mm, 2 2, 2.0—2.5 mm; taken from coarse coral sand and algal scrapings in | m of water, Rarotonga Is., Cook Islands. VOLUME 92, NUMBER 4 829 Fig. 11. Pendanthura rarotonga: a, Pereopod 1; b, Pereopod 2; c, Pereopod 7; d, Pleopod 1. 830 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Remarks.—The until-now monotypic genus Pendanthura was represent- ed by P. tanaiformis Menzies and Glynn from the Caribbean. The present species agrees closely with the generic diagnosis (Menzies and Glynn 1968:31) and with the type-species (see Kensley, in press, a) especially in the one-segmented mandibular palp, antennal structure, 3-segmented maxilliped with endite, short triangular carpus of the posterior pereopods and the short fused pleon. A few important differences easily separate the two species. These include more elongate posterior pereopods in P. rarotonga, a less developed lobe on the propodal palm of pereopod 1, more fringed spines on the inner face of the propodus of pereopod 1, a relatively longer pleon consisting of fused pleonites 1—S, pleonite 6 free, and the lack of strong pigmentation. Family Paranthuridae Paranthura astrolabium, new species Figs. 12-13 Description.—¢@: Integument not indurate, with scattered chromato- phores on cephalon, pereon, and pleon. Body proportions: C<1=2=3> 4 <5 >6> 7. Cephalon with very low rostral point; irregular band of pig- ment between eyes. Pleonites 1—5 fused, segments indicated laterally; pleon- ite 6 free, with deep middorsal incision in posterior margin. Telson elliptical/ oval. Antennular peduncle 4-segmented, segment 4 very short; flagellum of 2 articles, terminal article very short. Antennal peduncle 5-segmented, seg- ment 2 grooved to accommodate antennule; flagellum of single shortened and somewhat flattened setose article. Mandibular palp 3-segmented, ter- minal segment with 7 spines. Maxilla with about 10 distal serrations. Max- illipedal segment 2 with short triangular extension at inner distal angle; segment 3 elongate; 2 distal segments very short, setose. Pereopod 1 unguis of dactylus with oblique suture; propodus expanded, palm with evenly con- vex ridge on inner surface plus row of 14-16 setae; outer margin of palm Straight, with short proximal lobe. Pereopods 4—7 with propodus having 3 serrate sensory spines on posterior margin; carpus roughly triangular, with 2 serrate sensory spines on posterior margin. Pleopod 1 exopod operculi- form, with several elongate plumose setae on distal margin; endopod two- thirds length and about one-tenth width of exopod, with 3 distal plumose setae. Uropodal exopod ovoid, reaching beyond end of basis, distal margin serrate, apically acute, with elongate simple setae; endopod reaching slightly beyond telsonic apex, broadly ovate, with elongate simple setae. Etymology.—The specific name, proposed as a noun, is derived from Great Astrolabe Barrier Reef in the Fiji Islands. Material.—Holotype, USNM 172283, °, 4.5 mm; Paratype, USNM VOLUME 92, NUMBER 4 831 e Fig. 12. Paranthura astrolabium: a, Female in dorsal view; b, Antennule; c, Antenna; d, Maxilliped; e, Maxilla; f, Mandible; g, Pereopod 1. 172284, 2, 4.0 mm; Great Astrolabe Barrier Reef, off Ndravuni Is., Fiji Islands; taken from calcareous algal rubble in 5—13 m of water. Remarks.—Paranthura astrolabium differs very obviously from P. po- lynesica (described below) from the Cook Islands in size, chromatophore pattern, telsonic and uropodal shape, length and relative proportions of the 832 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 13. Paranthura astrolabium: a, Pereopod 7; b, Pleopod 1; c, Uropodal endopod and basis; d, Uropodal exopod. VOLUME 92, NUMBER 4 833 endopod of pleopod 1, and setation/spination of pereopod | propodus. There is some similarity to P. lifuensis Stebbing, 1900, recorded from the Loyalty Islands, especially in telsonic shape and structure of pereopod 1. The en- dopod of pleopod 1 appears more elongate and setose in P. lifuensis, while the sixth pleonite apparently lacks the deep middorsal incision in the pos- terior margin as seen in P. astrolabium. In this latter feature there is a strong resemblance to P. ostergaardi Miller and Menzies, 1952, from Ha- wail, but this species has a distally rounded uropodal exopod (acute/dentate in P. astrolabium), a more setose palm of pereopod 1, and a more elongate endopod of pleopod 1. Paranthura polynesica, new species Figs. 14-15 Description.—°?: Integument not indurate, with regular arrangement of large chromatophores dorsally. Body proportions: C >1>2=3=4> 5 > 6> 7. Pleonites 1-5 fused, 6 free, with deep middorsal incision in pos- terior margin. Telson elongate/rectangular, distally truncate, margin with elongate setae. Antennular peduncle 4-segmented, segment 4 short; flagel- lum of 3 articles, 2 distal articles very short. Antennal peduncle 5-segment- ed, segment 2 grooved to accommodate antennule; flagellum of 1 (?2) arti- cles, slightly flattened and setose. Mandibular palp 3-segmented, middle segment twice length of segment 1; terminal segment with 6 fringed spines. Maxilla slender, with 8—10 serrations. Maxilliped 3-segmented, terminal seg- ment elongate, tapering; no tiny distal segments visible. Pereopod 1 prop- odus expanded, inner palmar margin evenly convex, with 3 proximal fringed spines, outer margin straight, with low triangular proximal process. Pereopods 2 and 3 subchelate, propodus not as expanded as pereopod 1, propodal palm with 5 sensory spines. Pereopods 4—7, propodus with 2 strong sensory spines on posterior margin; carpus roughly rectangular, with sen- sory spine at posterodistal corner. Pleopod 1 exopod operculiform, with several plumose setae on distal margin; endopod about half length of exo- pod, elongate/triangular, with 2 short terminal setae. Uropodal exopod reaching to endopod, elongate/oval, outer margin dentate, distally acute; endopod almost circular, with numerous elongate simple setae, just reaching beyond telsonic apex. Etymology.—The specific name is derived from Polynesia, the general descriptive term for the area in which the entire collection was made. Material.—Holotype, USNM 172285, 2°, 2.8 mm, Paratypes, USNM 172286, 3 2, 2.0-2.8 mm; Rarotonga Is., Cook Islands; taken from rubble on back reef in 1 m of water; Paratypes, USNM 172287, 2, 3.1 mm; Mangaia Is., Cook Islands; taken from algal carpet growing on limestone in about 0.5 m of water. 834 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON iq Fig. 14. Paranthura polynesica: a, Female in dorsal view; b, Mandible; c, Maxilliped; d, Antennule; e, Antenna; f, Pereopod 1; g, Pereopod 2. 4 Remarks.—Paranthura is perhaps the genus most requiring revision amongst all the Anthuridea, and to add two more names to an already daunting list may be regarded as irresponsible. It is necessary, however, to record the presence of species of Paranthura, especially from an area rarely collected. To give a new specific name, rather than to hide the material erroneously under an earlier name, or under the non-commital Paranthura Sp., 1S preferable. VOLUME 92, NUMBER 4 835 HN / j ‘\ Fig. 15. Paranthura polynesica: a, Telson; b, Uropod; c, Pleopod 1; d, Pereopod 7. Paranthura polynesica to some extent resembles P. bellicauda Miller and Menzies, 1952, from Hawaii, especially in the truncate telson and dentate uropodal exopod. P. bellicauda, however, possesses a much shorter pleon, and a more spinose/setose pereopod 1, and is a larger species (6 mm). Dif- 836 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON ferences are also seen in the uropodal endopod and in the relatively broader telson. Acknowledgments My thanks are due to Mr. C. A. Child of the Department of Invertebrate Zoology, Smithsonian Institution, for collecting the material described herein, and for his detailed field data: also to Dr. Thomas E. Bowman of the Smithsonian Institution, for reading the MS and making useful sugges- tions and criticisms. Literature Cited Kensley, B. 1978. The South African Museum’s Meiring Naude cruises Part 8. Isopoda Anthuridea.—Annals of the South African Museum 77: 1-235. . Inpress, a. The Anthuridea of Carrie Bow Cay, Belize (Crustacea, Isopoda).—Smith- sonian Contributions to Marine Sciences. . In press, b. Anthuridean Isopod Crustaceans from the International Indian Ocean Expedition 1960-65, in the Smithsonian Institution’s collections.—Smithsonian Contri- butions to Zoology. Menzies, R. J., and P. W. Glynn. 1968. The common marine isopod Crustacea of Puerto Rico.—Studies on the Fauna of Curacao and other Caribbean Islands 27:1-133. Miller, M. A., and R. J. Menzies. 1952. The isopod Crustacea of the Hawaiian Islands, III. Superfamily Flabellifera, Family Anthuridae.—Occasional Papers of the Bernice P. Bishop Museum, Honolulu, Hawaii. 21:1-15. Stebbing, T. R. R. 1900. On Crustacea brought by Dr Willey from the South Seas.—Willey’s Zoological Results 5:618—625. Department of Invertebrate Zoology, Smithsonian Institution, Washing- ton, D.C. 20560. PROC. BIOL. SOC. WASH. 92(4), 1979, pp. 837-862 THE HYDROPS YCHE SCALARIS GROUP IN VIRGINIA, WITH THE DESCRIPTION OF TWO NEW SPECIES (TRICHOPTERA: HYDROPSYCHIDAE) Oliver S. Flint, Jr., J. Reese Voshell, Jr., and Charles R. Parker Abstract.—Fifteen species of the Hydropsyche scalaris group are record- ed from Virginia. Four sites on the North and South Anna Rivers, which harbor 8 species, are characterized and physico-chemical data given. H. incommoda Hagen is redescribed and diagnosed on the basis of the type and recently collected material. H. incommoda, sensu Ross 1944, is de- scribed and named H. rossi n. sp., and H. bassi n. sp. is described. The ranges and habitats of the species are discussed and suggestions explaining the diversity of the fauna advanced. The Commonwealth of Virginia occupies a favorable position on the east- ern seaboard of the United States from the standpoint of its potential faunal diversity. The state cuts across 5 physiographic provinces, 4 life zones (sen- su Merriam) or 7 ‘“‘regions’’ (sensu Hoffman, 1969), and ranges in elevation from sea level to 1,743 m (5,720 feet). As a consequence, a considerable faunal diversity is to be expected, but, for most insect groups, not yet reported. Our efforts to investigate the aquatic fauna of the state, both on a broad survey basis and an in-depth analysis of certain sites, are now showing the hoped-for richness of fauna. As a result of an in-depth study of 4 sites on the North and South Anna Rivers by Voshell and Parker, 8 species of the Hydropsyche scalaris group were discovered. Attempts to identify these species soon convinced Flint that the taxonomy of the group badly needed study. Types of problem species were borrowed, and their study revealed that the trouble was based on a misidentification of the species H. incommoda Hagen with its subse- quent ramifications. This paper, then, is an attempt to explain the diversity of the fauna at the Anna Rivers sites, to rectify the systematic problems, and to present the distribution and zoogeography of the sca/aris group of species in the state. DESCRIPTION OF NORTH AND SOUTH ANNA RIVERS The North and South Anna Rivers are tributaries of the York River Basin (Map 1) which lies in the central and eastern section of Virginia. The York River Basin drains 6,892 km?; from the headwaters in Orange County to the mouth in Chesapeake Bay is a distance of approximately 193 km. The basin 838 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON varies in width from 8 km at the mouth of the York River to 64 km in the headwaters. The western part of the York River Basin lies within the Pied- mont Physiographic Province, and the eastern part lies within the Coastal Plain Province. The Piedmont exhibits a rolling surface of gentle slopes and mild relief. Elevation varies from over 549 m in the extreme western portion to 76 m at the eastern edge of the Piedmont. The Fall Line separates the Piedmont from the Coastal Plain. The Coastal Plain slopes gently eastward from the Fall Line to the Chesapeake Bay. Elevation varies from 76 m to 0 m at the mouth of the York River. The topography of the Coastal Plain is gently sloping hills and flat farmland. Most of the basin is forested (70%) or in cropland and pasture (22%); very little of the land area is classified as urban (2%). There is very little variation in climate within the York River Basin. The climate is moderate with an average annual temperature of 14°C; extremes from less than — 18°C to above 38°C are recorded. Average annual precipitation in the basin is approximately 109 cm, varying very little from 107-117 cm. Average annual snowfall is light, ranging from 25 cm along the Coastal Plain to 38 cm in the upper Piedmont (Virginia Division of Water Resources, 1970). The length of the course of the North Anna River (NAR) in the Piedmont is 91 km. The gradient averages 2.7 m/km along this course, with an alti- tudinal range from 264 to 18 m. The Piedmont section of the South Anna River (SAR) extends for 154 km, with an average gradient of 2.9 m/km and an altitudinal range from 458 to 18 m. Four sampling sites were established in the lower Piedmont for the purpose of analyzing the downstream effects of Lake Anna on the NAR by comparison with the free-flowing SAR. One Station was established on the NAR at Rt. 601, approximately 0.5 km below Lake Anna (Map 1, number 1). At this point the NAR serves as the bound- ary between Louisa and Spotsylvania Counties. Another station was estab- lished on the NAR approximately 32 km downstream at the Fall Line (Map 1, number 2). At this point the NAR serves as the boundary between Han- over and Caroline Counties. On the SAR, stations were established at Rt. 522 in Louisa County (Map 1, number 3) and approximately 54 km down- stream at the Fall Line (Map 1, number 4). The latter station at the Fall Line was on Rt. 657 in Hanover County. Both rivers meander slowly across the Piedmont. The elevation of the study area ranges from 55 to 20 m on the NAR and from 70 to 38 m on the SAR. The average slope of each river between the study sites is 1.1 m/km for the NAR and 0.6 m/km for the SAR. Within the study area the maximum slopes for each river (21.8 m/km for the NAR and 3.4 m/km for the SAR) occur at the Fall Line where the lower sampling sites were established. The average mean daily discharge of the NAR during the study was 12 m?/sec (range: 1—260 m/sec); the average for the SAR was 7 m*/sec (range: 2—77 m/sec). The rivers vary in width from 20 to 75 m along their courses. The NAR is a Sth order stream at both study 839 ‘(ge ‘d 998) soqis UOI}D9TIOO JBANY VUUY YINOS ‘p 3? ¢ ‘(gEg “d 92S) SOUS UOTDETIOD JOATY PUUY YWON ‘7 2 | ‘setdeds dnois syyjvos YY JO uoNnNquysiq— | dew 4WyMs IWWSId NVMOHO NINYSHSW SNONVOY a 430334, NOLS70H NOLS70H ce a Gs! ie yt ae SORT RTT FERC RETR 77 CTE Ty (QA CR UI RRS RR onf we, '4'S A'N DON gen mary j Cpe Ae ZUSTAS Se alae ee ent WINNIE i | aX | pope SE FOE G i] j' \ mma Lo J 24 Ps 7 ISSOY AHOASdONGAH V VGOWWOON] SHOASKOMGAH ISILLV4I AHOASdOYNGAH @ SOlIN OF Le pee fam teen as $49;@Wol}y OF ° WINIDUIA VOLUME 92, NUMBER 4 840 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Table 1.—Basic physical and chemical properties of the North and South Anna Rivers from June 1977 to June 1978. The upper number is the mean and the lower numbers in parentheses are the range. North Anna River South Anna River Parameter Rt. 601 Fall line Rt. 522 Fall line Temperature (°C) 18.4 18.7 16.7 17.6 (3.0-31.0) (2.5—29.0) (2.0—26.0) (4.0—28.0) Dissolved oxygen (ppm) 9.4 9.8 Sell 9.5 (6.0—13.5) (6.0—13.5) (6.0—13.0) (6.5—13.0) Dissolved oxygen saturation 0.99 1.05 0.93 1.00 (0.67-1.21) (0.79-1.27) (0.68-1.14) (0.78—1.20) Total alkalinity (ppm CaCO,) 18 18 25 26 (12-25) (12-29) (13-45) (15-35) Hydrogen ion concentration (pH) 6.86 7.00 6.80 7207, (6.20-7.60) (5.90-7.70) (6.30-7.35) (6.40-8.05) Specific conductance (umho/cm) 61 61 67 69 (40-92) (40-93) (20-125) (12-120) Orthophosphate (ppm) 0.05 0.03 0.13 0.08 (0.00-0.34) (0.00-0.06) (0.00-0.60) (0.00—0.40) Total phosphate (ppm) 0.48 0.52 0.51 0.51 (0.00—2.94) (0.00-3.56) (0.02-3.03) (0.07—2.64) Nitrate nitrogen (ppm) 0.362 0.615 0.480 0.443 Ammonia nitrogen (ppm) Sulfate (ppm) (0.000—0.804) (0.000—3.420) (0.249-0.848) (0.127—-0.707) 0.095 0.042 0.188 0.018 (0.000—0.238) (0.000—0.096) (0.000-0.516) (0.000—0.954) 11 (9-13) 11 (8-15) Bt (6-30) 1] (5-19) sites; the SAR is 4th order at the upper site and Sth order at the lower site. Both rivers are characterized by long stretches of slow-moving water with shifting-sand bottom, occasionally interspersed with riffles, logs, and ac- companying leaf debris. The substrate in the riffles where the quantitative benthic samples were collected consists primarily of cobble (64—256 mm). Two aquatic vascular plants are abundant in the SAR, Podostemum cera- tophyllum (river weed) and Justicia americana (water willow), whereas, only Justicia americana is present in the NAR. Riparian vegetation of both rivers is dominated by Betula nigra (river birch), Alnus serrulata (speckled alder), Platanus occidentalis (sycamore), Fraxinus americana (white ash), and Ilex decidua (possum hawthorn). Basic physical and chemical properties of the water at each of the sam- pling sites are presented in Table 1. The NAR and SAR are warm-water VOLUME 92, NUMBER 4 841 rivers of soft to medium hardness and circum-neutral pH, typical of the Piedmont. Dissolved oxygen concentration is usually near saturation in rif- fles. There are some differences in nutrient concentrations between the two rivers, probably resulting from the impoundment of the NAR. However, neither the NAR nor the SAR seems to exhibit any chemical properties that would be limiting factors for aquatic insect life, and, overall, both rivers appear ‘“‘healthy.”’ METHODS Larvae, pupae, and adults were collected. Larvae and pupae were col- lected in riffles with a Portable Invertebrate Box Sampler (PIBS) (Ellis- Rutter Assoc., Douglassville, Pa.), by dip-netting, and by handpicking. From June 1977 to June 1978, collections were made biweekly during the warmer months (April—October) and monthly in the colder months. The quantitative samples taken with the PIBS were preserved in the field. Basic physical and chemical parameters were measured concurrent with the ben- thic collections. Some mature larvae and pupae were kept alive and returned to the laboratory for rearing. Damp burlap, placed in 8 oz. glass jars and kept cool with a small amount of ice, proved very effective for keeping larvae and pupae alive. The glass jars were placed in styrofoam coolers designed to hold six canned beverages. Specimens were identified to species as reared adults or as pupae using the metamorphotype method. Adults were collected with portable black light traps and with lights (either black light or Coleman lantern) at white sheets of cloth. Adults were collected at various stations from spring through autumn; most stations had at least one adult collection in spring, summer, and fall in order to collect all of the species present at each station. The collecting traps or lights were usually set up at dusk and operated for 1 hour. Types of all species treated in this paper, plus certain other closely related species of the scalaris group, were borrowed from either the Illinois Natural History Survey (INHS) or the Museum of Comparative Zoology (MCZ). Identifications were made based on comparison with the types. In certain instances there are small differences between the type and Virginia material, or between examples from different parts of the range of the species. How- ever, these differences do not seem to be of specific value. All material listed in detail, and not specifically marked otherwise is in the collection of the National Museum of Natural History (USNM). Most of the material from Virginia is contained either in the collection of the Virginia Polytechnic Institute and State University (VPI&SU) or the USNM. 842 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON TAXONOMY Hydropsyche alvata Denning Figs. 1-5, Map 2 Hydropsyche alvata Denning, 1949, p. 40.—Gordon and Wallace, 1975, pp. 413, 415. Unzicker et al., 1970, p. 171. This poorly known species was described from Mississippi, Illinois and Michigan, and subsequently recorded from Georgia and Arkansas. It thus appears to be a basically southern species extending northward in the Mis- sissippi basin and along the Atlantic Coastal Plain and Piedmont. It is very closely related to H. orris and overlaps the latter in most char- acteristics. Apparent specific differences are to be found in the apex of the phallus which is clearly larger than the adjacent phallobase and has a domed mesal cavity in lateral aspect. In ventral aspect the inner margins of the lateral plates are sinuous and a bit more open posteriad than in orris. The figures here presented were prepared from a paratype from Momence, IIli- nois. Virginia records.—Hanover Co.: North Anna River at the Falls, 1 mile west of U.S. Rt. 1, 21 June 1977, Parker, 3 6, VPI&SU, 1 6, USNM. Louisa Co.: South Anna River at Rt. 522, 22 June 1977, Voshell, 2 6, VPI&SU; same, 5 July 1977, 1 6. Hydropsyche bassi Flint, Voshell, and Parker, n. sp. Figs. 19-23, Map 5 Hydropsyche scalaris (nec Hagen): Schuster & Etnier, 1978, p. 89 (material from Tennessee).—Etnier & Schuster, 1979, p. 16. This species is very close to scalaris from which it differs in being smaller, in minor genitalic differences, and especially in the size of the eyes of the males. In scalaris the eyes are only slightly enlarged (in frontal aspect the eye is about % the interocular distance), but in bassi the eyes are huge, being distinctly wider than the frontal interocular distance. The genitalia of the two species are virtually identical, with the apex of the apical segment of the clasper which is truncate in bassi rather than pointed (much like Figures 2 or 8) being the most distinctive. The tip of the phallus is also proportionately shorter in both lateral and ventral aspects in bassi. Adult.—Length of forewing, 9 mm. Color brown in alcohol; forewing strongly mottled. Eye of male in frontal aspect wider than interocular dis- tance (5:4). Male genitalia: Ninth segment with anterior margin rounded, and enlarged lateroventrally; with a low dorsomesal crest. Tenth tergite with apex truncate in lateral aspect, with a large setiferous wart and a group of setae dorsally; in dorsal aspect with apex truncate and narrowly excised mesally. Clasper with basal segment long, slender, and straight; apical seg- ment in lateral aspect with tip curved into a dorsally directed point, in caudal VOLUME 92, NUMBER 4 843 Figs. 1-13. AHydropsyche alvata Denning: 1, male genitalia, lateral; 2, clasper, ventral; 3, ninth and tenth terga, dorsal; 4, tip of phallus, ventral; 5, tip of phallus, lateral. H. orris Ross: 6, tip of phallus, lateral; 7, tip of phallus, ventral. H. incommoda Hagen: 8, clasper, ventral; 9, tip of phallus (type), ventral; 10, tip of phallus, lateral; 11, male genitalia, lateral; 12, ninth and tenth terga, dorsal; 13, tip of phallus (Virginia specimen), ventral. 844 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON oS ve i | is j ,A \ Fa SY NYA A i 3 Ne? SY X S| wy YD ay ‘) Wy --=<5.2 7 ~-(Z - uA) 23 Hydropsyche rossi n. sp.: 14, male genitalia, lateral; 15, clasper, ventral; 16, Figs. 14-23. ninth and tenth terga, dorsal; 17, tip of phallus, lateral; 18, tip of phallus, ventral. H. bassi n. 19, tip of phallus, lateral; 20, tip of phallus, ventral; 21, clasper, ventral; 22, male Sp.: genitalia, lateral; 23, ninth and tenth terga, dorsal. VOLUME 92, NUMBER 4 845 aspect slightly bowed and squarely truncate. Phallus tubular, phallobase slightly constricted before apex; in lateral aspect with ventral margin rapidly tapering dorsad to a blunt apex, mesal dome distinctly elevated, lateral flange up-turned laterally; in ventral aspect the inner margin of lateral flange almost straight, mesal cavity deep and semicircular. Material.—Holotype, male: Virginia, Russell Co., Big Cedar Creek at route 19, May 1978, Michael Bass. USNM Type 76302. Paratypes: Tennes- see, Knox Co., Beaver Creek at Pellissippi Pkwy., just south of Oak Ridge, 9 May 1979, Etnier et al., 1 6; same, 18 April 1976, Etnier, 1 d metamor- photype (Univ. Tennessee). Etymology.—We dedicate this species to the collector of the type, Dr. Michael L. Bass, Mary Washington College. Hydropsyche catawba Ross Map 2 Hydropsyche catawba Ross, 1939, p. 67.—Gordon & Wallace, 1975, pp. 413, 416. , This rarely reported species appears to be limited to Piedmont rivers. It has been reported from Georgia and North Carolina and now Virginia. Virginia records.—Hanover Co.: North Anna River at the Falls, 1 mile west of U.S. Rt. 1, 2 June 1977, Parker, 2 d, VPI&SU; same, 12 June 1978, Flint, 2 6; same, 21 June 1977, Parker, 4 6, VPI&SU, 2 6, USNM; South Anna River at Rt. 657, 11 September 1978, Voshell, 2 6, VPI&SU. Louisa Co.: North Anna River at Rt. 601, 12 June 1978, Flint, 1 d 1 @. Hydropsyche dicantha Ross Map 3 Hydropsyche dicantha Ross, 1938b, p. 146; 1944, p. 102.—Morse & Bickle, 1953, p. 71.—Etnier, 1965, p. 146.—Longridge & Hilsenhoff, 1973, p. 176.—Roy & Harper, 1975, p. 1082.—Schuster & Etnier, 1978, p. 82.— Etnier & Schuster, 1979, p. 16. This species has a wide range in northeastern North America, apparently reaching its southern limits in Virginia and Tennessee. It is recorded from the District of Columbia, Kentucky, Michigan, Minnesota, New Hampshire, New York, Ontario, Quebec, Tennessee, Wisconsin, and there are examples in the USNM from Ohio and Pennsylvania. Virginia records.—Shenandoah Co.: Elizabeth Furnace, 25 May 1966, Beard, 1 1-22 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 846 SolIW Ob $19;9WO}] OS ‘soiseds dnois supjpos *Y Jo uoynqmsiq— Zz dep NVMOH9 NINYSH3aW SHONVOU.u oiige LS EO ee SS aa TET SI ERS oe HON 173M0d way BOTS ae Gj iN} iN ota et LK j aid aa Sy nS : 5 pice ae SISNAlddISSISSIN SHOASdOYGAH ¥ VEMVLVO AHOASdOMGAH VLVAIV SHOASdOYNGAH @ vig th iS \ yi S ¢ ‘ ‘ f ? oi Cx | VOLUME 92, NUMBER 4 847 Hydropsyche fattigi Ross Map | Hydropsyche fattigi Ross, 1941, pp. 88-89.—Gordon & Wallace, 1975, pp. 411, 415.—Etnier & Schuster, 1979, p. 16. This species has been reported from Georgia and Tennessee previously. It is widely distributed in the Piedmont of Virginia where it reaches the northernmost known limits of its distribution. Virginia records.—Campbell Co.: Bennett Farm, about 28.3 km SE of Rustburg on Rt. 615, 27 June 1978, Zimmerman, | 6, VPI&SU. Fauquier Co.: Broad Run, Thorofare Gap, 26 July 1975, Flint, 17 3 32 2; same, 20 September 1965, 3 ¢ 1 2. Hanover Co.: South Anna River at Rt. 657, 12 June 1978, Voshell, 10 ¢, VPI&SU; same, Flint, 2 ¢d 2 2, USNM; same, 20 June 1977, Parker, 1 6, VPI&SU; same, 21 June 1977, Voshell, 13 3 5 2; same, 19 July 1977, 2 56; same, 3 August 1977, 1 ¢6; same, 11 September 1978, 5 5; same, 13 September 1977, Parker, 2 6. Louisa Co.: South Anna River at Rt. 522, 19 July 1977, Voshell, 3 3d, VPI&SU. Aydropsyche hageni Banks Map 4 Hydropsyche hageni Banks, 1905, p. 14.—Denning, 1943, pp. 119-—120.— Ross, 1944, p. 173.—Longridge & Hilsenhoff, 1975, p. 176.—Etnier & Schuster, 1979, p. 16. H. hageni has a wide distribution throughout the eastern United States, being reported from Illinois, Kentucky, Manitoba, Maryland, Minnesota, North Carolina, Tennessee, and Wisconsin. In addition, we have seen ma- terial from the District of Columbia. Virginia records.—Fairfax Co., Giles Co., Montgomery Co., Page Co., Rockingham Co., Shenandoah Co., Warren Co., Wythe Co. Range of dates, 3 May to 30 August. Hydropsyche hoffmani Ross Map 4 Hydropsyche hoffmani Ross, 1962, pp. 129-130.—Schuster & Etnier, 1978, pp. 98-100. This species, which has not been reported previously outside of Virginia, is widespread in Virginia and also has been taken in Maryland on the banks of the Potomac River. Virginia records.—Arlington Co., Fairfax Co., Montgomery Co., Page Co., Roanoke Co., Rockingham Co., Shenandoah Co., Warren Co. Range of dates, 17 May to 10 September. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 848 SOIIN OF 7 $10;9WOI}} OS toy dS AWWSIG Rechte NINYSHSW ‘so1sods dnois supjpos * JO uolNgIysIq—'¢ dey ONO 33033d , NOLS10H NOLS10H d's ‘M4'N HONITO SIYNVIVIS SHOASdOUGAH ¥ VOINTIVHLdO SHOASdOUGAH VHLNVOIG SHOASdOUGAH @ VOLUME 92, NUMBER 4 849 Hydropsyche incommoda Hagen Figs. 8-13, Map 1 Hydropsyche incommoda Hagen, 1861, p. 290.—Ross, 1938a, p. 17.—Fi- scher, 1963, p. 43. Hydropsyche orris (nec Ross): Wallace, 1974, pp. 549-550.—Gordon & Wallace, 1975, pp. 405-423.— Wallace & Malas, 1976, p. 208.—Wallace et al., 1977, pp. 506-532. Because incommoda Hagen is one of the oldest names for a North Amer- ican species of Hydropsyche, it has been mentioned in the literature many times (Fischer, 1963). However, it is doubtful if it has ever been correctly applied with the exception of Ross’ (1938a) designation and illustration of the lectotype. Although Ross (1944) stabilized the identification and nomen- clature in the scalaris group, he unfortunately misidentified incommoda (he apparently did not have any examples of the true incommoda available at that time), and as a result the name has been consistently misapplied since then. True incommoda appears to be most closely related to orris Ross and alvata Denning, and much more distantly to the species long known as incommoda. In recent years with work on Atlantic Coastal Plain and Pied- mont rivers the species has been rediscovered, but apparently consistently misreported as orris Ross. We have not been able to find any examples of the true orris from east of the Appalachians. The three species alvata, incommoda, and orris, all agree in having the apex by the tenth tergum rather rounded in lateral aspect, the apical segment of the clasper in caudal aspect tapering to a sharp point, and the lateral lobes of the phallus broadly rounded in lateral aspect. The 3 species may be distinguished by the shape of apex of the phallus, especially in ventral view, by the following triplet: 1. Lateral plates of phallus in ventral view with inner margins of lateral flanges virtually parallel, mesal cavity deep ......... (Figs. 6-7) orris 2. Lateral plates of phallus in ventral view with inner margins of lateral flanges divergent and slightly sinuous, mesal cavity shallow and widely Opened: posteniad ......0. s2o8... 6. (Figs. 8-13) incommoda 3. Lateral plates of phallus in ventral view with inner margins of lateral flanges very slightly divergent, and sinuous, mesal cavity deep and only slightly more opened posteriad than anteriad .. (Figs. 1-5) alvata Adult.—Length of forewing, 8.5—10 mm. Color brown; forewing strongly mottled, with a large pale spot at the anal angle. Eye of male in frontal aspect slightly more than half width of interocular distance (3:5). Male gen- italia: Ninth segment with anterior margin broadly rounded; in dorsal aspect with a mesal crest. Tenth tergite with apex rounded in lateral aspect, with a single large setiferous wart; in dorsal aspect with a variably-formed, shal- PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 850 “sotoeds dnois supjpos “Fy JO uoNNqmMsiIq—‘p dew NVMOH9 NINYSH3SW SMONVOU. NOLS10H H i TRIN Se RA RSS tO ag WoNITO xs ja Wes co zhhs A y IGYVWNO371 SHOASdOUGAH W INVW4JOH SHOASdOUGAH INSOVH SHOASdONGAH @ SOlIW OF $10j;9W0)|1 OS VINIDUIA VOLUME 92, NUMBER 4 851 low mesal excision. Clasper with basal segment long, slender and straight; apical segment in lateral aspect with tip curved into a dorsally directed point, in caudal aspect with tip tapered to a point. Phallus tubular, phallo- base slightly constricted before apex, in lateral aspect with ventral margin tapering dorsad to a broadly rounded apex, with mesal dome slightly ele- vated, in mesial aspect with inner margin of lateral flange slightly sin- uous, flared laterad at apex, with mesal cavity very shallow (in the type the lateral plates are crushed slightly with a resultant skewing of the parts). Material.—Lectotype, male [M.C.Z.]: Georgien, Winthem, Type 11018. Virginia records.—Campbell Co.: Ferris Farm, about 10 km SE of Rustburg on Rt. 615, 13 June 1978, Zimmerman, 2 6, VPI&SU; Bennett Farm, about 28 km SE of Rustburg on Rt. 615, 27 June 1978, Zimmerman, 22 6 8 °, VPI&SU; same, 4 July 1978, 1 6; same, 18 July 1978, 1 ¢d. Hanover Co.: North Anna River at the Falls, 1 mile west of U.S. Rt. 1, 27 May 1978, Voshell, 1 6, VPI&SU; South Anna River at Rt. 657, 8 October 1977, killed 5 May 1978, Voshell, 1 ¢ mmt, VPI&SU. Louisa Co.: North Anna River at Rt. 601, 12 June—14 September 1977-78, Voshell and Flint, many 66 and 22 and mmt, VPI&SU, USNM. New Kent Co.: Pamunkey R., at White House, 15 July 1971,;.Simmons, 4 ¢ 1 ¢@. In addition we have seen material of this species from Florida, Georgia, and Maryland. Hydropsyche leonardi Ross Map 4 Hydropsyche leonardi Ross, 1938b, pp. 145—146.—Schuster & Etnier, 1978, pp. 100-102. This seldom encountered species has been reported only from Michigan and Virginia. It seems to be found in riffles in larger rivers that are clean and fast-flowing. Virginia records.—Page Co.: South Fork of the Shenandoah River at Grove Hill, 12 April-9 June 1972, Surber, 1 ¢, VPI&SU. Roanoke Co.: Roanoke River at Rt. 419 in Salem, 5 July 1975, Etnier, 1 d mmt, same, 10 July 1975, Schuster, 7 mmt. Shenandoah Co.: Shenandoah River near Woodstock, 20 October 1962, Flint, 1 ¢. Hydropsyche mississippiensis Flint Map 2 Hydropsyche mississippiensis Flint, 1972, p. 80.—Schuster & Etnier, 1978, pp. 95—96.—Etnier & Schuster, 1979, p. 16. Although previously known only from Mississippi and Tennessee, we have records from Louisiana and South Carolina in addition to Virginia. The species seems to be primarily an inhabitant of Piedmont rivers of the southeastern United States. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 852 ‘saiseds dnois supjpos ‘HY Jo uonNnquysIq—¢ dey NVMOHO NIMUAHAN = AMONVOU.4 NOLS 10H dWYMS IWWSIG oe DLS" M4N HONITO 7 . = i \ i aN ISSV@ AHOASdOUGAH ¥ ~SIYVINNSA SHOASdOUGAH VLVYSIVHd SHOASKdOYGAH @ SOuIN OF 4 x Z ; Ss. BY f ; x ; Ne ; Hf W/ i S1pjoWO]Iy OS ae ( e is p J ‘ 2 , = SS K / i VOLUME 92, NUMBER 4 853 Virginia records.—Bedford Co.: Carter Mill Creek, Rt. 734, 16 July 1975, Flint & Ronderos, 4 d 1 2. Louisa Co.: North Anna River at Rt. 601, 12 June 1978, Voshell & Flint, 2 6, VPI&SU, 3 6, USNM; same, 21 June 1977, Voshell, 3 d, VPI&SU; same, 20 July 1977, 1 ¢; same, 30 July 1977, Parker, 1 d; same, 14 September 1977, 3 6; South Anna River at Rt. 522, aeuly 1977, Voshell, 13°0,) VPP & SU: Hydropsyche opthalmica Flint Map 3 Hydropsyche opthalmica Flint, 1965, p. 169. This species, unreported since its original description from West Virginia, is now recorded from several localities in Virginia. The records all cluster around rather large streams that are clear, cool and fast flowing. Virginia records.—Augusta Co.: Middle River at Mt. Meridian, 20 May 1978, Seagle, 1 d, VPI&SU. Bath Co.: Jackson River, approx. 400 m below Back Creek, 1973, Strickler, unpublished M.S. Thesis, VPI&SU. Page Co.: South Fork of the Shenandoah River at Brumback’s Cabin, 10 May 1971, Surber, 3 d6, VPI&SU, 1 6, USNM. Rockingham Co.: South Fork of the Shenandoah River at Goods Mill, 26 May 1°71, Surber, 1 ¢, VPI&SU. Shenandoah Co.: North Fork of the Shenandoah River, Rt. 707, 15 June 1971, Surber, 1 6, VPI&SU; North Fork of the Shenandoah River, Swope Hollow, Rt. 661, Mauertown, 21 July 1971, 1 ¢; North Fork of the Shen- andoah River, 1971, Surber, 39 ¢6, VPI&SU. Warren Co.: South Fork of the Shenandoah River, Bentonville Bridge, 24 May 1971, Surber, 1 6, VPI&SU, 1 6, USNM; South Fork of the Shenandoah River, Public Boat Landing at Front Royal, 30 May 1978, Seagle, 3 6, VPI&SU; same, 8 June 1977, Parker, 2 d 4 2; same, 27 June 1978, Seagle, 2 6. Hydropsyche phalerata Hagen Map 5 Hydropsyche phalerata Hagen, 1861, p. 287.—Ross, 1944, p. 102.—Leonard & Leonard, 1949, p. 10.—Fischer, 1963, p. 77.—Etnier, 1965, p. 146.— Longridge & Hilsenhoff, 1973, p. 176.—Resh, 1975, p. 12.—Schuster & Etnier, 1978, pp. 78—80.—Etnier & Schuster, 1979, p. 16. This is a commonly collected species, widespread in the eastern half of the United States and southeastern Canada. Records are available from the District of Columbia, Georgia, Illinois, Indiana, Kansas, Kentucky, Mich- igan, Minnesota, New Jersey, New York, North Carolina, Ohio, Pennsy!l- vania, Tennessee, Wisconsin, and Canada. Material of the species from Florida and Maryland is also present in the USNM. Virginia records.—Arlington Co., Bedford Co., Carroll Co., Fairfax Co.., 854 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Floyd Co., Giies Co., Hanover Co., Louisa Co., Montgomery Co., Rich- mond City, Rockingham Co., Shenandoah Co., Warren Co. Range of dates, 25 May to 27 September. Hydropsyche rossi Flint, Voshell, and Parker, n. sp. Figs. 14-18, Map 1 Hydropsyche incommoda (nec Hagen): Ross, 1944, p. 106.—Unzicker et al., 1970, p. 171.—Gordon & Wallace, 1975, pp. 405—423.—Resh, 1975, p. 12.—Wallace, et al., 1977, pp. 506-532.—Schuster & Etnier, 1979, pp. 92—93.—Etnier & Schuster, 1979, p. 16. This is a rather common southern species, ranging as far north at least as Illinois in the Mississippi Basin and Virginia on the Atlantic Coast. This species is the one misidentified by Ross in 1944 as incommoda, and reported as such by all subsequent authors. Based on this concept, the species has been recorded from Arkansas, Florida, Georgia, Illinois, Kentucky, Loui- siana, North Carolina, Tennessee, and we have records from Mississippi, Missouri, South Carolina, and Virginia. The species differs from alvata, incommoda, and orris on the basis of the more rectangular apex of the tenth tergum, the more nearly truncate apex of the clasper, and in the apex of the phallus. In rossi the lateral plates are thin and long in lateral aspect, and are very narrow and widely open in ventral aspect. There is some variation in the width of the lateral flange, some examples from Illinois being almost as wide as Fig. 13. Adult.—Length of forewing, 10-11 mm. Color brown; forewing strongly mottled, generally with a paler band longitudinally in middle. Eye of male in frontal aspect almost as wide as interocular distance (4:5.5). Male geni- talia: Ninth segment with anterior margin broadly rounded; in dorsal aspect with a mesal crest. Tenth tergite with apex truncate in lateral aspect, with a large setiferous wart and a row of setae dorsally; in dorsal aspect with apex truncate and narrow excised mesally. Clasper with basal segment long, slender, and straight; apical segment in lateral aspect with tip curved into a dorsally directed point, in caudal aspect with tip obliquely truncate. Phal- lus tubular, phallobase scarcely constricted before apex, in lateral aspect with ventral margin rapidly tapering dorsad to a long, narrow apex, with mesal dome slightly elevated; in ventral aspect with inner margin of lateral flange very narrow, with scarcely any mesal cavity, widely open apicad. Material.—Holotype, male: MISSISSIPPI, Forrest Co., Hattiesburg, 25 Sept. 1967, Bryant Mather. USNM Type 76303. Paratypes: Same, 1 <6; same, 19 Sept. 1967, 1 6 3 2; Warren Co., Bovina, 21-28 April 1971, Mather, | ¢6 1 2; Washington Co., Wayside, 27 June 1966, Bruce, | d; State College, 16 June 1972, Robinson, 1 ¢ 1 2; same, 25-30 Apr. 1966, Davis, 3 6 20 2, 1.N.H.S.; Marshall Co, 1 mi. W. Waterford, 26 June 1973, VOLUME 92, NUMBER 4 855 Stark, 7 ¢ 3 2; Stoneville, May 1959, Pfrimmer, 4 3, I.N.H.S.; Leland, 17-28 May 1958, Pfrimmer, 1 3, I.N.H.S.; Columbus, Tombigbee R., 21 May 1976, Schuster, 1 6, U.T. ARKANSAS, Bradley-Drew County line, Saline R., 21 May 1977, Etnier et al.,2 ¢, U.T. FLORIDA, Alachua Co., Gainesville, 26 Apr. 1967, Wirth, 1 ¢; Columbia Co., O’Leno State Park, 31 May 1966, Beard, 1 ¢ 2 2; Marion Co., Juniper Springs, 28 Apr. 1970, Wirth, 6 6 5 2; Lake Co., Tavares, 23 Mar. 1936, Young, 4 6 5 2, I.N.H.S.; Jackson Co., Blue Springs Cr., 3 mi. E. Marianna, 5 June 1940, Berner, 1 6 4 2, I.N.H-S. GEORGIA, Effingham Co., Savanna R. at end Rt. 275, 15 May 1970, Wal- lace & Sherberger, 5 ¢; Altamaha R., nr. Baxley, Hatch Power Plant, 22 Apr. 1974, Ga. Power Co., 2 ¢ 1 @. ILLINOIS, Carmi, Littke Wabash R., 24 Apr. 1935, Frison, 1 6, I.N.H.S.; Carlyle, 9 Aug. 1943, Sanderson & Leighton, 3 ¢ 1 °,1.N.H.S.; Mahomet, 3 Aug. 1937, Ross & Burks, 12 ¢ 10 2, I.N.H.S.; Oakwood, 20 Sept. 1935, DeLong & Ross, 1 6, I.N.H.S.; Benton, 11 June 1945, Ross, 16 1 2,I1.N.H.S.; Monticello, 7 May 1936, Ross & Burks, 1 6, I.N.H.S.; Pontiac, 22 Aug. 1938, Ross, 3 6, 1.N.H.S.; Kappa, Mackinaw R., 14 Sept. 1937, Ross & Burks 4 6, I.N.H.S.; Momence, 16 Aug. 1938, Ross & Burks, 4 6, 1.N.H.S.; Momence, Kankakee R., 29 May 1939, Frison & Ross, 1 6,1.N.H.S.; Poplar Bluff, 2 June 1948, Frison, 1 ¢, I.N.H.S.; Urbana, 1- 7 July 1958, Klatt, 19 d 2 29, 1.N.H.S.; same, 22 July 1958; 11 6 1 2; same, 6 Aug. 1958, 2 6 22 2; same, 20 Aug. 1958, 52 6; same, 26 Aug. 1958, 7 o. KENTUCKY, 25 mi. N.E. Bowling Green, 3 Sept. 1967, Druckenbrod, 1 6. LOUISIANA, Bossier Parish, 10 May 1938, Turner, 1 ¢ 1 2; Curtis, Red R. Valley Exp. Sta., 22 July 1958, 2 6 2 2, I.N.H.S.; Washington Parish, Silver Cr. 6 mi N.E. Franklinton, 5 May 1979, Louton, | ¢, U.T. MISSOURI, Columbia, 21 July 1974, Craig, 1 ¢6. NORTH CAROLINA, Statesville, Catawba R., 23 Apr. 1938, Ross & Burks, 1 6, I.N.H.S. SOUTH CAROLINA, Jackson, 22 Apr. 1969, Tarpley, 2 ¢; Savanna R., Calhoun Falls (Ca. 3 mi. S. of town), 27 Apr. 1969, Wallace, 3 d6; Colleton Co., Hwy, 61, Edisto R., 5 mi. E. Canadys, 19-22 May 1968, Cartwright, 26 1°. TENNESSEE, Madison Co., Jackson, 13 May 1957, 2 6, I.N.H.S.; same, 20 May 1957, 1 2°; Lauderdale Co.; Ripley, 6 May 1957, 1 6, 1.N.H.S.; Cumberland Co., Crossville, 13 Sept. 1970, 1 6, U.T.; Rob- ertson Co., 1 June 1966, 1 6, U.T.; Fentress Co., 12-14 July 1972, Van Landingham, 1 6, U.T.; same 20-24 July 1972, 1 6. TEXAS, Brazos Co., Little Brazos R., at Tex. 21, 23 May 1977, 13 6, U.T. VIRGINIA, Louisa Co., North Anna River at Rt. 601, 17 December 1977, emerged 5 March 1978, Voshell, 1 6, VPI&SU; same, emerged 12 Apnil 1978, Parker, 1 ¢; same, 14 September 1977, Voshell, 3 d, 1 2 mmt, VPI&SU; Pittsylvania Co., Chatham, 7 July 1963, Tarpley, 1 6. Etymology.—We dedicate this species to the late Dr. H. H. Ross, in 856 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON memory of his many contributions to trichopterology, especially the basic work on the scalaris group. Hydropsyche scalaris Hagen Map 3 Hydropsyche scalaris Hagen, 1861, p. 286.—Ross, 1944, p. 106.—Blickle & Morse, 1966, p. 6.—Nimmo, 1966, p. 691.—Unzicker et al., 1970, p. 171.—Edwards, 1973, p. 504.—Schuster & Etnier, 1978, pp. 87-90 (in part). Although this species is widespread throughout eastern North America, records before Ross (1944) are suspect because of widespread misidentifi- cations. There appear to be valid records from Arkansas, Georgia, Indiana, Maine, Minnesota, Missouri, Oklahoma, Ontario, Quebec, Texas, Wiscon- sin, and material at the USNM from Maryland and New York. Virginia records.—Augusta Co., Bath Co., Bedford Co., Botetourt Co., Carroll Co., Craig Co., Floyd Co., Giles Co., Montgomery Co., Rockbridge Co., Rockingham Co., Shenandoah Co., Washington Co. Range of dates, 21 May to 20 October. Hydropsyche venularis Banks Map 5 Hydropsyche venularis Banks, 1914, p. 252.—Resh, 1975, p. 12.—Wallace, 1975, pp. 463—472.—Schuster & Etnier, 1978, pp. 96-98. This is often a common species in the southeastern United States. It is found from the Coastal Plain into the Cumberland Plateau, generally favor- ing medium to large rivers. Apparently valid records are from the District of Columbia, Georgia, Kentucky, North Carolina, Tennessee, and in ma- terial at the USNM, South Carolina. Virginia records.—Bedford Co., Caroline Co., Carroll Co., Fauquier Co., Floyd Co., Giles Co., Hanover Co.. Louisa Co., Montgomery Co., Rich- mond City. Range of dates, 3 May to 28 September. DISTRIBUTION We have found 15 species belonging to the H. scalaris group in Virginia. This is approximately the same number reported in two other southeastern States: 14 in Tennessee (Etnier and Schuster, 1979) and 11 in Georgia (Gor- don and Wallace, 1975). Fewer species have been reported in midwestern and northern United States. Ross (1944) reported 11 species belonging to the H. scalaris group from Illinois in his comprehensive survey of the state fauna. Etnier (1965) included 10 species of this group in his list of Minnesota VOLUME 92, NUMBER 4 857 Trichoptera, Longridge and Hilsenhoff (1973) listed 9 species from Wiscon- sin, Leonard and Leonard (1949) recorded 9 species from Michigan, and Resh (1975) substantiated 8 species in Kentucky. Morse and Blickle (1953) only recorded 1 species in New Hampshire, and in 1966 (Blickle & Morse) listed a different species from Maine. Nimmo (1966) and Roy and Harper (1975) listed 5 species from Quebec, however, some of these species were misidentified. Of the 15 species of the H. scalaris group that occur in Virginia (Maps 1-5), 6 are southeastern species that appear to reach their northern limit in Virginia, or in two instances, Maryland or the District of Columbia. These species are H. alvata, H. catawba, H. fattigi, H. incommoda, H. missis- sippiensis, and H. venularis. Two species have northern distributions that extend into the southeast. H. leonardi is known only from Michigan and Virginia, and in Virginia is limited to the Ridge and Valley Physiographic Province. H. dicantha is known from 7 northern states and Canada and extends into Kentucky, Ten- nessee, and Virginia. Within Virginia it has been collected in the northern mountains and across the Potomac in the District of Columbia along the Fall Line. Thus, at least 8 species of the H. scalaris group that occur in Virginia appear to be southern species reaching their northern limit or north- ern species reaching their southern limit. In addition, 3 species are known only from Virginia and one bordering state. H. hoffmani has been recorded from Maryland, H. opthalmica has been recorded from West Virginia, and H. bassi has been recorded from Tennessee. The 4 remaining species, H. hageni, H. phalerata, H. rossi, and H. scalaris are widespread in the east. Of the 15 species in the H. scalaris group found in Virginia, 12 species have been collected in the Piedmont, but only 9 species have been collected at higher elevations to the west of the Piedmont (Table 2). In Georgia, Gordon and Wallace (1975) found 7 species in the Piedmont and 6 species in the headwaters. Combining the records from Georgia and Virginia, 14 species belonging to the H. scalaris group have been found in the Piedmont and 13 species at higher elevations to the west. In Virginia we have collected 8 species from the lower Piedmont sections of the NAR and SAR in the York River Basin. Gordon and Wallace (1975) reported only 5 species from the lower Piedmont sections of the Savannah River Basin. A review of the distribution records of the species in the H. scalaris group raises the ques- tions: why are there so many species in the southeastern Piedmont, and, in particular, why are there so many species in a relatively small area of the lower Piedmont in Virginia? Ross (1963) speculated that the entire scalaris complex of the H. scalaris group evolved in the temperate deciduous forest, in large creeks or rivers with sufficient current to result in rock or gravel bottom. The number of 858 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Table 2.—Distribution of species belonging to the H. scalaris group in Virginia and Georgia. Georgia records are from Gordon and Wallace (1975). Virginia Georgia—Savannah River Basin Higher Lower elevations Piedmont west of sections of Lower Species Piedmont Piedmont NAR and SAR Headwaters Piedmont Piedmont H. alvata X X xX H. bassi X H. bidens xX xX xX H. catawba X X xX H. demora xX H. dicantha X xX H. fattigi X xX X H. hageni xX xX H. hoffmani X xX H. incommoda xX xX xX xX H. leonardi X H. mississippiensis xX xX H. opthalmica xX H. phalerata XxX XxX X H. rossi X xX X xX H. scalaris X xX xX X xX HZ. nr. scalaris X XxX H. venularis X xX xX xX xX different species found in southeastern Piedmont rivers provides support for Ross’ hypothesis. The following comparison of the results of several recent ecological investigations with our observations from the NAR and SAR is an attempt to further explain the distribution of this abundant group of caddisflies. Cummins (1977) stated that greater diversity of species is expected in the midreaches (width: 10-75 m; stream order: 4—6) than in the upstream or downriver areas of a River Continuum. His delimitation of midreaches co- incides with the widths and stream orders of the NAR and SAR. He men- tioned thermal diversity, variety of organic substrates, and physical heter- Ogeneity as important factors affecting the diversity of the midreaches. In regard to thermal regime, Sweeney and Vannote (1978) hypothesized that the distribution of species, both locally and over large geographic areas, is limited by lowered fecundity as adult size gradually diminishes in streams of increasingly cold or warm temperature cycles. Each species has an op- timum temperature regime for larval growth that permits the insect to achieve maximum adult weight and fecundity. Small adults and reduced fecundity result when temperatures are either warmed or cooled with re- spect to more optimal thermal conditions. However, aquatic insects have VOLUME 92, NUMBER 4 859 evolved means of surviving unfavorable conditions such as diapause, hi- bernation, estivation, or quiescence, all of which can occur in various stages. Therefore, it is possible to have species with different optimum temperature regimes coexisting in the same habitat if the annual temperature cycle of the habitat includes the optimum temperature of all the species. In fact, one would expect higher species diversity in habitats with more diverse temperature regimes. Vannote (in Cummins, 1977) has suggested that great- er diversity of species is expected in the midreaches because the water temperature is more variable than in the groundwater controlled headwaters or in the large volume rivers. These hypotheses certainly appear to be sup- ported by the high diversity of species occurring in the low Piedmont sec- tions of the NAR and SAR, where the temperature varies as much as 28°C at a given site during the year (Table 1). The wide range in the annual temperature cycle allows the coexistence of both spring and summer species. In addition, the wide range in temperatures is conducive to range extensions by northern and southern species. Boreal species are able to reduce the competition for resources by dispersing downstream into the Piedmont. Austral species are able to accomplish the same thing by dis- persing upstream to-the limits of their optimum thermal regime. H. dicantha is a good example of a boreal species which is found in both the mountains and Piedmont in Virginia (Map 3). H. alvata, H. catawba, H. mississip- piensis, H. fattigi, H. incommoda, and H. rossi are examples of austral species that occur in the Piedmont of Virginia, particularly the lower Pied- mont. The diversity of species in the Piedmont may also be explained by the overall physical heterogeneity of rivers in this physiographic province. For example, the most abundant size-class of substrate in individual benthic samples from the NAR and SAR varied from boulder (>256 mm) to coarse sand (0.5—1 mm). The flow measured at individual benthic samples varied from 168 cm/sec to <1 cm/sec. The variable flow and substrate reflect the irregular longitudinal profile of Piedmont rivers. While much of the NAR has only a moderate gradient, the gradient at the Fall Line (21.8 m/km) is equivalent to that found in many headwater streams. Because riparian vege- tation does not entirely shade Piedmont rivers, aquatic vascular plants are sometimes abundant. Podostemum, a preferred attachment site for many Hydropsychidae, was found in mats as dense as 698.7 g dry weight/m? in the SAR. Snags are another characteristic feature of Piedmont rivers. As river banks erode, large trees topple in and remain in the river for many years. Such snags have been shown to be preferred attachment sites for some Hydropsychidae that are most common in very large rivers (Wallace et al., 1977). The available food cannot be overlooked as an explanation for the diver- sity of species belonging to the H. scalaris group in the Piedmont. Gordon 860 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON and Wallace (1975) indicated that subtle differences in feeding habits of the larvae and change in available food along the course of the stream affect the distribution of Hydropsychidae. Wallace et al. (1977) stated that the type and size of drifting stream seston has influenced the evolution of filter- feeding Trichoptera. Species that occur in fast-flowing headwater streams have capture nets with large mesh sizes, and species that inhabit slow-flow- ing coastal rivers have capture nets with very small mesh openings. The evolution of the filter-feeding Trichoptera that occupy both habitats has produced individual species capable of cropping various types and sizes of seston. Cummins (1977) reported that the ratio of course particulate organic matter (CPOM) to fine particulate organic matter (FPOM) decreases in the midreaches because large amounts of FPOM are introduced from tributaries and sloughed from algal mats and vascular plant beds. However, there are still inputs of CPOM in the midreaches from both riparian vegetation and aquatic vascular plants. In addition, the flood plain of the Piedmont is a source of input of CPOM and FPOM. Therefore, the variety of types and sizes of seston in Piedmont rivers is probably reflected in the number of different species in the H. scalaris group that occur there. It does not appear that the seston input from the reservoir on the NAR affects the diversity of species in the H. scalaris group. All species except H. rossi have been collected from both the regulated NAR and free-flowing SAR. ACKNOWLEDGMENTS We wish to acknowledge the willing cooperation of Ms. M. K. Thayer, Museum of Comparative Zoology, Harvard Univ., and Dr. J. D. Unzicker, Illinois Natural History Survey for making available the types and other valuable material in their institutions’ collections. Dr. D. A. Etnier, Univer- sity of Tennessee, Dr. P. P. Harper, Université de Montréal, and Dr. J. B. Wallace, University of Georgia submitted material for reidentification from their collections. Mr. B. C. Kondratieff, VPI&SU, assisted in the field and laboratory. The base map of Virginia was kindly provided by Dr. R. E. Jenkins, Virginia Commonwealth University. It is a modification of the U.S.G.S. State of Virginia, scale 1:500,000 map, 1957 edition, and was compiled by N. M. Burkhead, D. J. Jenkins, and R. E. Jenkins, with financial support from the Virginia Commission of Game and Inland Fisheries. LITERATURE CITED Banks, N. 1905. Descriptions of new Nearctic neuropteroid insects.—Trans. Amer. Ent. Soc. 32:1-20. . 1914. American Trichoptera—notes and descriptions.—Can. Ent. 46:252-258. Blickle, R. L. and W. J. Morse. 1966. The caddisflies (Trichoptera) of Maine, excepting the family Hydroptilidae.—Maine Agr. Exp. Sta. Tech. Bull. 24:1-12. VOLUME 92, NUMBER 4 861 Cummins, K. W. 1977. From headwater streams to rivers.—Amer. Biol. Teacher 39:305—312. Denning, D. G. 1943. The Hydropsychidae of Minnesota (Trichoptera).—Entom. Amer. 23:101-127. . 1949. New species of Nearctic caddis flies.—Bull. Brooklyn Ent. Soc. 44:37-48. Edwards, S. W. 1973. Texas caddisflies —Texas J. Sci. 24:491-516. Etnier, D. A. 1965. An annotated list of the Trichoptera of Minnesota, with description of a new species.—Ent. News 76:141-152. Etnier, D. A. and G. A. Schuster. 1979. An annotated list of Trichoptera (caddisflies) of Tennessee.—J. Tennessee Acad. Sci. 54:15—22. Fischer, F. C. J. 1963. Hydropsychidae, Arctopsychidae.—Trichopt. Catalog IV: 1-225. Flint, O. S., Jr. 1965. New species of Trichoptera from the United States.—Proc. Ent. Soc. Washington 67: 168-176. . 1972. Three new caddisflies from the southeastern United States.—J. Georgia Ent. Soc. 7:79-82. Gordon, A. E. and J. B. Wallace. 1975. Distribution of the Hydropsychidae (Trichoptera) in the Savannah River Basin of North Carolina, South Carolina, and Georgia.—Hydro- biologia 46:405—423. Hagen, H. H. 1861. Synopsis of the Neuroptera of North America.—Smiths. Misc. Coll. 4(1):xx + 1-347. Hoffman, R. L. 1969. The biotic regions of Virginia.— Virginia Polytech. Inst., Res. Div. Bull. 48:23-62. Leonard, J. W. and F. A. Leonard. 1949. An annotated list of Michigan Trichoptera.—Oc- casional Papers Mus. Zool. Univ. Michigan 522, 35 pp. Longridge, J. L. and W. L. Hilsenhoff. 1973. Annotated list of Trichoptera (caddisflies) in Wisconsin.—Wisconsin Acad. Arts Sci. & Lett. 61:173-183. Morse, W. J. and R. L. Blickle. 1953. A checklist of the Trichoptera (Caddis Flies) of New Hampshire.—Ent. News 64:68-102. Nimmo, A. P. 1966. A list of the Trichoptera taken at Montreal and Chambly, Quebec, with descriptions of three new species.—Can. Ent. 98:688-693. Resh, V. H. 1975. A distributional study of the caddisflies of Kentucky.—Trans. Kentucky Acad. Sci. 36:6—16. Ross, H. H. 1938a. Lectotypes of North American caddisflies in the Museum of Comparative Zoology.—Psyche 45:1-61. . 1938b. Descriptions of Nearctic caddis flies (Trichoptera).—Bull. Ill. Nat. Hist. Surv. 21:101-183. . 1939. New species of Trichoptera from the Appalachian Region.—Proc. Ent. Soc. Washington 41:65-—72. . 1941. Descriptions and records of North American Trichoptera.—Trans. Amer. Ent. Soc. 67:35-126. . 1944. The caddis flies, or Trichoptera, of Illinois.—Bull. Ill. Nat. Hist. Surv. 23:1- 326. . 1962. Three new species of Trichoptera from eastern North America.—Ent. News 73:129-133. . 1963. Stream communities and terrestrial biomes.—Arch. Hydrobiol. 59:235—242. Roy, D. and P. P. Harper. 1975. Nouvelles mentions de trichopteres du Quebec et description de Limnephilus nimmoi sp. nov. (Limnephilidae).—Can. J. Zool. 53:1080-1088. Schuster, G. A. and D. A. Etnier. 1978. A manual for the identification of the larvae of the caddisfly genera Hydropsyche Pictet and Symphitopsyche Ulmer in eastern and central North America (Trichoptera: Hydropsychidae).—USEPA, Environmental Monitoring and Support Laboratory, EPA-600/4-78-060, pp. xii + 129. Sweeney, B. W. and R. L. Vannote. 1978. Size variation and the distribution of hemi- metabo!lous aquatic insects: two thermal equilibrium hypotheses.—Science 200:444-446. 862 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Unzicker, J. D., L. Aggus, and L. O. Warren. 1970. A preliminary list of the Arkansas Trichoptera.—J. Georgia Ent. Soc. 5:167—174. Virginia Division of Water Resources. 1970. York River Basin: Comprehensive Water Re- sources Plan. Volume I—Introduction.—Planning Bulletin 225, 112 pp. Wallace, J. B. 1974. Silk spinning as an escape mechanism in Hydropsyche orris larvae following removal from water (Trichoptera: Hydropsychidae).—Ann. Ent. Soc. Amer. 68:549-550. . 1975. Food partitioning in net-spinning Trichoptera larvae: Hydropsyche venularis, Cheumatopsyche etrona, and Macronema zebratum (Hydropsychidae).—Ann. Ent. Soc. Amer. 68:463—472. Wallace, J. B. and D. Malas. 1976. The significance of the elongate, rectangular mesh found in capture nets of fine particle feeding Trichoptera larvae.—Arch. Hydrobiol. 77:205- DAWe Wallace, J. B., J. R. Webster, and W. R. Woodall. 1977. The role of filter feeders in flowing waters.—Arch. Hydrobiol. 79:506—532. (OSF) Smithsonian Institution, Washington, D.C. 20560; (JRV and CRP) Dept. of Entomology, Virginia Polytechnic Institute and State University, Blacksburg, VA 24061. PROC. BIOL. SOC. WASH. 92(4), 1979, pp. 863-872 A REDESCRIPTION OF THE WOOD-RASPING AMPHIPOD TROPICHELURA GOMEZI ORTIZ, 1976 (CHELURIDAE) FROM THE FLORIDA KEYS, WITH NOTES ON ITS DISTRIBUTION AND ECOLOGY James Darwin Thomas Abstract.—The wood-rasping amphipod, Tropichelura gomezi Ortiz, 1976, is redescribed from the Florida Keys to include diagnostic characters omitted in the original description. Morphological characters and distribu- tional patterns of 7. gomezi are compared with those of the other member of the genus, Tropichelura insulae (Calman, 1910). Laboratory studies show that T. gomezi defends its burrow entrance from other members of its species, but tolerates the presence of limnoriid isopods with which it co- occurs. A lectotype for Tropichelura insulae is designated. Introduction Recent investigations on the littoral marine amphipod fauna of the Florida Keys have yielded a wood-rasping amphipod, tentatively identified as Tropichelura gomezi Ortiz, 1976, originally described from Cuban waters. Adult specimens of 7. gomezi from the Florida Keys exhibit distinctive sexual and morphological differences not included in Ortiz’ original descrip- tion. Ortiz’ collection from which the holotype (a female, 5.0 mm in body length) was designated lacked larger specimens in the 8-12 mm size range where primary sexual characters are more pronounced. Females in the 4— 5 mm range superficially resemble males but are readily distinguished from them by the presence of paired telsonic setae. Although requests for topotypes of 7. gomezi have been unsuccessful, examination of identical material of wood-rasping amphipods from Belize, Puerto Rico, and the Tortugas tends to confirm material from the Florida Keys as T. gomezi. On this assumption, this study purports to redescribe and refigure Tropichelura gomezi and provide new information on its ecology and life history. T. gomezi is compared with the only other member of the genus, T. insulae (Calman, 1910). The Cheluridae, a small family of corophioidean amphipods, was revised by Barnard (1959) to include 3 genera: Chelura Philippi, Nippochelura Bar- nard, and Tropichelura Barnard. Chelura, represented by a single species, 864 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Chelura terebrans Philippi, 1839, is widely distributed in warm temperate seas and is distinguished by the presence of an inner ramus on uropod 3. Nippochelura, also represented by a single species, Nippochelura brevi- cauda (Shiino, 1948), is reported only from Japanese waters and is charac- terized by having uropods 2 and 3 uniramous. Tropichelura is tropical in distribution and composed of 2 species, Tropichelura insulae (Calman, 1910), and Tropichelura gomezi Ortiz, 1976 and is distinguished by a biramous second uropod and a uniramous third uropod. The ecology of chelurids dates from 1839, when Philippi first described Chelura terebrans from marine wood borings. Because of the economic importance of chelurids, the literature on this small group of marine am- phipods is surprisingly extensive; more than 65 papers on C. terebrans alone have been published. Investigations on the ecology of C. terebrans by Bar- nard (1955) determined that limnoriid isopods initiated burrows that were subsequently invaded and enlarged by Chelura. Miller (1924) reported Tropichelura insulae abundant in marine timbers in Samoa, where this species enlarged the burrows of Paralimnoria andrewsi (Calman) for its own occupancy. Ortiz (1976) recorded 7. gomezi associated with the isopod Limnoria platycauda (Menzies) from the Gulf of Batabanao, Cuba. In the Florida Keys, 7. gomezi was taken with Paralimnoria andrewsi, Limnoria platycauda, and Limnoria simulata (Menzies). My observations show that T. gomezi is a very active burrower, connecting and enlarging limnoriid burrows into large, unroofed galleries by rasping and furrowing of the softer wood grain. Tropichelura gomezi is stenohaline and occurs in protected coastal bays and other shallow habitats (less than 3 m) that are not subject to significant salinity fluctuations. T. gomezi was conspicuously absent from harbor and open ocean situations, even though numerous samples of submerged timbers were examined from these areas. Samples taken from harbor areas held large numbers of limnoriids, while the ocean samples were riddled with the shipworm, Toredo. Tropichelura gomezi Ortiz Figs. 1-4 Tropichelura gomezi Ortiz, 1976: pp. 21-26, figs. 1-2. Diagnosis.—Antenna 1, article 1 expanded anterolaterally, articles 1-3 cylindrical, length 2.5 width. Maxilla 2, internal lobe with 14—16 terminal setae. Gnathopod 2, palm transverse, defined by strong tooth, 90% of pro- podus. Pereopod 7, article 2 linear, length 2 width. Description.—Male, 12.0 mm. Head normal for genus; supra-antennal line present; eyes large, black-pigmented. Antenna | short, accessory flagellum small, 1-segmented; flagellum short, VOLUME 92, NUMBER 4 865 Fig. 1. Tropichelura gomezi: a, Lateral view, 12.0 mm ¢; b, Antenna 2, 2: c, Gnatho- pod, @. sparsely setose distally. Antenna 2 stout; flagellum of 1 clavate article: sparsely setose distally. Upper lip with epistome produced; labrum flattened distally. Mandible with 3-segmented palp, article 2 with row of oblique setae, article 3 with numerous comb setae, 2 apical setae; molar large, molar surface rasp-like; incisor toothed; 8 raker spines present. Lower lip with inner lobes unde- veloped; mandibular process present. Maxilla 1 with 2-segmented palp, 2nd segment with 5 spines and 2 apical setae; outer plate with 9 pectinate spines; inner plate reduced, 6 plumose setae medially. Maxilla 2, inner plate with 16-17 plumose setae; outer plate with 10-12 plumose setae. Maxilliped, palp 4-segmented, exceeding outer plate; terminal article with distal spine. Coxae 1-7 small; 5-6 bilobed; gills sac-like, present on coxae 2-6; Gnathopod | large; palm transverse, defined by tooth. Gnathopod 2 mi- nutely subchelate; articles 2-5 with long, pectinate setae anteriorly; article 5 with 5S rows of comb spines. Pereopod 3, article 2 slightly expanded anteriorly, anterior margin with 13 stout setae, posterior margin bare; article 3 lacking marginal setae; article 7 with 2 accessory nails. Pereopod 4, article 2 slightly produced anteriorly, anterior margin with 10 stout setae, posterior margin bare; article 7 with 2 accessory nails. Pereopod 5, coxal lobes subequal; article 1 bilobed; article 2, widely expanded, posterior margin with 8 plumose setae; article 4, pos- terior margin bare; article 6 with 5 large spines laterally, paralleled by 5 thinner spines; 2 large spines present at junction of articles 6 and 7. Per- eopod 6, article 2 slightly expanded, posterior margin bearing 8 plumose PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 866 =, eR Uloe: d = 6) ys ey: 3 Fy \ \ XY ON cee, (a,b,d,e,9) Tropichelura gomezi, 3: a, Lower lip; b, Upper lip, c, d, Maxilla 1; e, Mandible; Fig. 2. f, Maxilliped; g, Maxilla 2. 867 VOLUME 92, NUMBER 4 \\¢ _@) Fig. 3. Tropichelura gomezi, 3: a, Pleopod 1; b, Pleopod 2; c, Pleopod 3; d, Uropod 2; e, Uropod 1. 868 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 4. Tropichelura gomezi: Telson and uropod 3 of ¢ and 9. setae; article 6 with 3 pairs oblique spines; article 7, plumose setae present. Pereopod 7, article 2, length twice width, posterior margin with 14 plumose setae; article 6 with 6 pairs of large, oblique spines; 2 large spines present at junction of articles 6 and 7; article 7, single plumose seta present. Pleon segment 3 with 3 dentate tubercles posterodorsally. Pleopod pe- duncles each with 2 coupling hooks and single plumose seta. Uropod 1 biramous, inner ramus longest; peduncle normal. Uropod 2 biramous, rami serrate; peduncle flattened and expanded, serrate laterally; few short setae present in serrations. Uropod 3, uniramous, ramus flattened and ovate, moderately serrate; few short setae between serrations. Telson apically serrate; notches with minute setae. Females.—Antenna 2, flagellum heavily setose distally. Gnathopod 2 smaller than but similar to male. Pereopods 6 and 7, posterior edge article 2 heavily setose in ovigerous specimens. Urosome with numerous, long setae laterally and ventrally. Telson with 4 apical notches, each bearing long, paired setae. Oostegites present on pereopods 2-S. Distribution.—Tropical Atlantic: Florida Keys, Dry Tortugas, Cuba, Belize, and Puerto Rico. Ecology.—Taken from sublittoral timbers and decaying red mangrove roots. Relationships.—While T. gomezi and T. insulae have obvious morpho- logical differences on the specific level (antenna 1, articles 1-3; palm, gnathopod 1; article 2, pereopod 7), they also share several similarities wor- 869 VOLUME 92, NUMBER 4 EEE EESS EE OS 9eJ9S LIOYS ‘S9Yyd}OU Z 9e19S [PUIWIS} p 9e}as JeUTUIID} sired p 9e19S WOYS ‘S9yd}OU Z uos]9} suo] yoys ‘osieds ae}as Zuo] avjas ‘asojas ATIABOY gejas ‘asojas A[IABOY yoys ‘asireds 9e}as ¢n-[n OpIM ‘oads “SIAO ‘gads ‘3IAO UI 9S0}aS se Suc] jsowye ‘ysod UI dSO}OS ATIABIY Z UIpIM Ajiavoy JOU Z “We ‘soyeu se popuedxe A[peoig Z “We ‘ye adpo “jsod ‘soyew se XZ YsUZ] ‘IeoUT] Z “We L ‘pd snpodoid snpodoid %0¢ “pouysp %06 *Y}00} SuoNs Aq poonpol ‘soyeul se Ayyeom ‘onbijgo wyed poonpol ‘soyeul se pouyop ‘asioasuen wyed | ‘us 9819S [CUIWIS} 9e}9S [eUIUIS} soyeul se p UUM 9qQoyT [eUuteIUI soyew se Q] YUM 9qQO] [euso}Ur 7 ‘xeul soyew Td 0S se yysua] *A]jeIsIp Td WET :0RI198 gsojos ATIAvOY WNTesey ‘gsojas ATIABVOY WNT[IseY gsojas ATIAvOY WINTfodey jeisip Moz YIM wNTTesey Z ‘ue Yip YIpIMm x ¢°7 x9 YIsUZ] ‘AT[VNUSA yisugy ‘jestIpuryAs SABNUOD ‘poua}ey fe 67 “*T “ye SAyesoqey] sojew se PAGO) Ge Coal peal soyeul se -olojue popuedxs | ‘We [ ‘que we ON ee (Ww ¢*¢ ‘oTeUI9z) (wu ¢°g ‘oyeu!) (wur ¢°9 ‘oTewoj) (Wu ¢‘Q] ‘oTeut) Ia}oRIeYyD aDjNSUl * apjnsul *], 1Z20UI08 *T, 1ZawW08 "TJ, 2s @ QU SS a Se eS ee eee ‘aDjNSU *[, pue IZaWOS *] JO SIOJOVIBYS ps}da]as JO suosuedwojg— | 9198 L 870 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON thy of mention. Females of 7. gomezi show a resemblance to males of T. insulae in the setose conditions of the flagellum on antenna 2, the urosome, and apical telsonic setae. Males of T. gomezi have sparsely setose urosomes as do females of 7. insulae. The relative amount of setae on the urosome is considered a diagnostic character and the two species of Tropichelura show a relationship in this condition; the males of 7. gomezi being similar to females of T. insulae, and females of T. gomezi resembling males of T. insulae. In the original species description, Ortiz (1976) reported that 7. gomezi differed from 7. insulae in having a quadrangular versus rectangular uro- some, and in the relation of the size of the third uropod. My observations show these characters to be variable and of questionable diagnostic use. Material examined:—INDIAN OCEAN: Tropichelura insulae (Calman, 1910).—British Museum (Natural History), Lectotype: 1978.291.1 (herein designated); male, 8.0 mm, taken from wooden pilings in Flying Fish Cove, Christmas Island, 1909. Paralectotypes: 1909.5.19.305-314. PACIFIC OCEAN: Tropichelura insulae.—National Museum Natural History, USNM 151391, Pearl Harbor, 1949. ATLANTIC OCEAN: Tropichelura gomezi Ortiz, 1976.—USNM 172091, Florida Keys, 1978.—USNM 1972096, Dry Tortugas, 1908.—JDT Bel. 2, Carrie Bow Cay, Belize, 1979.— Magueyes Island, Puerto Rico, specimens sent by Roger Zimmerman, 1978. Remarks.—Barnard (1959, 1971) reported 7. insulae as circumtropical and occurring throughout the Caribbean region. However, these records were no doubt erroneous and included records of the then undescribed T. gomezi. Distribution of 7. gom: zi in southern Florida and the Bahamian- Caribbean province is not known and awaits studies on additional material from these areas. Of pertinent interest would be material from the tropical coast of western Africa, a region from which no chelurids have been re- ported, although they are probably present there. Tropichelura is tropical in distritution, Nippochelura and Chelura are warm-temperate. 7. insulae is widespread in the Pacific and Indian Oceans, while 7. gomezi appears limited to the tropical Atlantic. Chelura terebrans ranges widely, being taken from both coasts of the U.S., Bermuda, western Europe, the Mediterranean, West and South Africa, the Suez Canal, and Australia. Nippochelura is endemic and recorded only from coastal Japan. Dispersal of chelurids is probably effected by timbers, ships, buoys, and other wooden flotsam. The present distribution of chelurids reflects some degree of isolation and speciation. As rafting is an effective dispersal mech- anism across large oceanic regions, chelurids might therefore be expected to occur widely in coastal situations throughout the world, especially where near-oceanic conditions persisted in close proximity to land. While this hy- pothesis holds true for the species of Tropichelura, Chelura is found in both estuarine and near-oceanic situations. VOLUME 92, NUMBER 4 871 The paucity of information in tropical areas probably reflects inadequate sampling. On the other hand, the voluminous published information per- taining to C. terebrans is a tribute to the negative economic impact of this wood-destroying organism. Laboratory investigations were conducted on 7. gomezi in submerged timbers, and showed the amphipod to move actively throughout the exca- vated galleries constantly probing burrow entrances and entering unoccu- pied areas. Large numbers of limnoriids were also present in the galleries, but were seemingly ignored by 7. gomezi; at least no apparent interaction between the two was detected. Defensive posturing behavior by T. gomezi was also noted. Thus, when a free-roaming individual investigated a burrow already occupied, the animal occupying the burrow would turn on its back and place the dorsal surface of its urosome (with uropods flared laterally) across the burrow entrance and thereby prevent further encroachment by the potential invader. The heavily calcified, quadrate urosome seems well-adapted to such defensive behavior. As 7. gomezi infests timbers in large numbers, overcrowding would presumably be lessened by such spatial partitioning of its burrows. Specimen size appéars to play little part in defensive posturing as large and small specimens seemed equally successful in repelling intruders. No studies were made on rates of wood-rasping in 7. gomezi and other chelurids; although presumably tropical forms, working in higher average temperatures would degrade wooden structures at a faster rate than their temperate counterparts. For further detailed discussion of chelurid ecology, the reader may consult the work of Barnard (1955) on the behavioral ecology of Chelura terebrans in Los Angeles Harbor. Acknowledgments I would like to thank Joan Ellis of the British Museum (Natural History), Dr. J. L. Barnard of the National Museum of Natural History, and Roger Zimmerman of the University of Puerto Rico for the loan of specimens during this study. Dr. Barnard, Dr. Pat McGlaughlin, and Dr. E. L. Bous- field kindly reviewed a preliminary manuscript. Dr. Bill Kruczynski iden- tified the limnoriid isopods taken during the study. Thanks are also due to Linda Lutz for inking the plates and preparing them for publication. This research was supported by the National Science Foundation, grant #DEB77-15883. Literature Cited Barnard, J. L. 1955. The wood-boring habits of Chelura terebrans Philippi in Los Angeles Harbor.—Essays Nat. Sci. Honor Capt. Allan Hancock, Hancock Fd. pp. 87-98, pls. 1-2. 872 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON . 1959. Generic partition in the amphipod family Cheluridae, marine wood borers.— Pacific Nat. 1(3):1-12, figs. 1-5. 1971. Keys to the Hawaiian marine Gammaridea, 0-30 meters.—Smith. Contr. Zo- ology 58:1—135, figs. 1-68. Calman, W. T. 1910. On two new species of wood-boring Crustacea from Christmas Island.— Ann. Mag. Nat. Hist. (8)5:181—186, pl. 5. Miller, R. C. 1924. Wood-boring Crustacea from Hawaii and Samoa.—Univ. Calif. Pub. Zool. 26(8): 159-164, pls. 12, 13. Philippi, A. 1839. Einige zoologische notizen.—Arch. Naturgesch. 5:113-134, pls. 3-4. Ortiz, M. 1976. Un neuvo antipodo perforador de madera (Amphipoda, Gammaridea, Che- luridae) de aguas Cubanas.—Invest. Marinas 8:21-26, figs. 1-2. Newfound Harbor Marine Institute, Big Pine Key, Florida 33043. PROC. BIOL. SOC. WASH. 92(4), 1979, pp. 873-875 THE CORRECT NAME OF THE HELIOPORAN OCTOCORAL LITHOTELESTO MICROPORA BAYER AND MUZIK Frederick M. Bayer Abstract.—Comparison of Recent Lithotelesto micropora Bayer & Mu- zik, 1977, with Cretaceous Primnoa gracilis Nielsen, 1925, shows that the two forms are congeneric. Synonymy of P. gracilis Nielsen, 1925, with Epiphaxum auloporoides Lonsdale, 1850, demonstrated by Voigt (1958), makes Lithotelesto Bayer & Muzik, 1977, a junior subjective synonym of Epiphaxum Lonsdale, 1850. Specimens of Epiphaxum are reported from the Indian Ocean south of Madagascar. Since publication of the description of Lithotelesto micropora Bayer and Muzik (1977:976), it has become clear that at least the genus already had been described as a fossil. Lonsdale (1850:261, pl. 18, figs. 35—37) described from the Chalk Formation of Sussex, England, a small, encrusting coral named Epiphaxum auloporoides having narrow, corrugated stolons con- taining at intervals ‘‘visceral cavities . . . provided with eight indentations or blunt lamellae.’’ As illustrated by Lonsdale, the stolons are very similar to those of Lithotelesto, but no specimens were available for direct com- parison. Somewhat later, Nielsen (1925:5, figs. 2-3) described from the Danian of Denmark a coral called Primnoa gracilis having a grooved calcareous axis grossly similar to that of certain gorgonaceans of the family Primnoidae but, unlike them, hollow and tubular rather than solid. Still later, Voigt (1958:9, pl. 1, figs. 1-7; 2, figs. 7-12; 10, fig. 3) demonstrated, on the basis of material from both lower and upper Maastrichtian from several localities in Ger- many, that the tubular ‘‘axes’’ of Primnoa gracilis Nielsen are, in fact, detached calicular tubes belonging to Epiphaxum. Comparison of the orig- inal material of Lithotelesto micropora with specimens of Primnoa gracilis from Denmark sent to the Smithsonian Institution by the Mineralogical and Geological Museum of the University of Copenhagen shows beyond doubt that the two are congeneric. As Primnoa gracilis has been demonstrated to be a synonym of Epiphaxum auloporoides Lonsdale (Voigt, 1958), the cor- rect generic name of Lithotelesto Bayer & Muzik, 1977, is Epiphaxum Lonsdale, 1850. 874 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Order HELIOPORACEA Bock, 1938 Coenothecalia Bourne, 1895:468. Helioporidea Bock, 1938:49. Helioporaria Stiasny, 1939:368. Helioporacea Bayer & Muzik, 1977:983 (nom. correct.). Family Lithotelestidae Bayer & Muzik, 1977 Article 40 of the International Code of Zoological Nomenclature requires that this family name be retained even though the name of the type-genus is now considered a junior synonym. So far as I can determine, no family- group name has ever been based on the generic name Epiphaxum. Voigt (1958) placed that genus in the family Clavulartidae, as I had done previously (Bayer, 1956:F'184). Genus Lpiphaxum Lonsdale, 1850 Epiphaxum Lonsdale, 1850:261 (type-species, Epiphaxum auloporoides Lonsdale, by monotypy). Primnoa.—Nielsen, 1925:5 (not Primnoa Lamouroux, 1812). Lithotelesto Bayer & Muzik, 1977:983 (type-species, Lithotelesto micro- pora Bayer & Muzik, by original designation and monotypy). Stratigraphic range.—Upper Cretaceous (Maastrichtian) to Recent. Geographic range.—Northwestern Europe (Upper Cretaceous); Lesser Antilles (Recent); Indian Ocean south of Madagascar (Recent). Epiphaxum auloporoides Lonsdale, 1850 Epiphaxum auloporoides Lonsdale, 1850:261, pl. 18, figs. 35—37.—Voigt, 1958:9, pl. 1, figs. 1-7; pl. 2, figs. 7-12; pl. 10, fig. 3. Primnoa gracilis Nielsen, 1925:5, figs. 2-3. Distribution.—Chalk Formation, Sussex (Lonsdale); Bryozoan Chalk at Faxe, Rejstrup and Herfggle, and the Chalk at Stevns Klint, Denmark (Niel- sen); Turonian, vicinity of Rostov, U.S.S.R., Campanian near Lagerdorf, Lower and Upper Maastrichtian from Hemmoor and several other localities in Western Germany (Voigt). Epiphaxum micropora (Bayer & Muzik, 1977), comb. nov. Lithotelesto micropora Bayer & Muzik, 1977:976, figs. 1-6. Distribution.—Lesser Antilles (Barbados). ?Indian Ocean (south of Mad- agascar). Remarks.—lIt is regrettable that Verrill made no mention of the locality and depth of his material in the explanation of his unpublished figures, which VOLUME 92, NUMBER 4 875 are the first known evidence of the occurrence of Epiphaxum in the Recent fauna (Bayer & Muzik, 1977). As those figures were included among the illustrations of the octocorals collected by the Blake, his specimens almost certainly were West Indian and may even have come from Barbados, as the Blake made 29 bottom stations there 5—10 March 1879. Specimens obtained from sediment samples collected at Walter Shoal south of Madagascar (33°12.0’S, 43°58.2’E, 360-200 m, Marion Dufresne cruise MD 08, sta. 6, DC 3, 16 March 1976) were correctly identified as Lithotelesto by Dr. H. Zibrowius and sent to me for verification. Unfortu- nately, those specimens retain no trace of soft tissue, making it impossible to compare anatomical and spicular characters with those of Epiphaxum micropora. AS some calices show development of eight distinct skeletal septa, which so far have not been observed in West Indian material, I am not able to state with certainty that this material is specifically identical with E. micropora. Acknowledgments I am indebted to Dr. Porter M. Kier of the Department of Paleobiology, National Museum of Natural History, for valuable advice in the paleonto- logical interpretation of Lithotelesto and Epiphaxum, and to Dr. Thomas E. Bowman, Department of Invertebrate Zoology, for critical reading of the manuscript. Literature Cited Bayer, Frederick M. 1956. Octocorallia. Jz Raymond C. Moore (Ed.), Treatise on Invertebrate Paleontology, part F:166—231.—Geological Society of America and University of Kansas Press, xx + 498 pp. Bayer, Frederick M., and Katherine M. Muzik. 1977. An Atlantic helioporan coral (Coelen- terata: Octocorallia).—Proc. Biol. Soc. Wash. 90(4):975—-984, figs. 1-6. Bock, Sixten. 1938. The alcyonarian genus Bathyalcyon.—Kungl. Svenska Vet. Akademiens Handlingar (3)16(5):1—54, pls. 1-2. Bourne, Gilbert C. 1895. On the structure and affinities of Heliopora coerulea, Pallas. With some observations on the structure of Xenia and Heteroxenia.—Phil. Trans. Roy. Soc. London 25:205-—243, figs. 1-22. Lonsdale, William. 1850. Descriptions of the fossils of the Chalk Formation. Notes on the corals. Jn Frederick Dixon, The geology and fossils of the Tertiary and Cretaceous formations of Sussex.—London, Longman, Brown, Green, and Longmans. Pp. 237- 324, pls. 18, 18A. Nielsen, K. Briinnich. 1925. Nogle nye Octocoraller fra Danienet.—Medd. fra Dansk Geol. Forening, Kgbenhavn 6(28): 1-6, figs. 1-3. Stiasny, Gustav. 1939. System der Octocorallia.—Zool. Meded. Leiden 21:367-368. Voigt, Ehrhard. 1958. Untersuchungen an Oktokorallen aus der oberen Kreide.—Mitt. Geol. Staatsinst. Hamburg 27:5-49, figs. 1-8, pls. 1-13. Department of Invertebrate Zoology, National Museum of Natural His- tory, Smithsonian Institution, Washington, D.C. 20560. PROC. BIOL. SOC. WASH. 92(4), 1979, pp. 876-882 DISTICHOGORGIA SCONSA, A NEW GENUS AND SPECIES OF CHRYSOGORGIID OCTOCORAL (COELENTERATA: ANTHOZOA) FROM THE BLAKE PLATEAU OFF NORTHERN FLORIDA Frederick M. Bayer Abstract.—Distichogorgia sconsa, a new genus and species of chryso- gorgiid octocoral from the Blake Plateau off Northern Florida is described and is figured by means of drawings and scanning electron micrographs. The genus and species are compared with those genera and species to which they are most similar, and the genus is incorporated in a new key to genera of the family Chrysogorgiidae. In the course of revisionary investigations of the octocoral fauna of the western Atlantic, based on the extensive collections obtained by the re- search vessels Gerda, Gilliss, Columbus Iselin, and John Elliott Pillsbury of the University of Miami, together with older collections in the National Museum of Natural History, Smithsonian Institution, many novelties have come to light. Although most of these will be published in the revisionary work proper, some are of sufficient interest to warrant publication in ad- vance of the larger work in order to bring them to the attention of interested investigators who may be in a position to increase our knowledge of them. Among these is a chrysogorgiid apparently allied to the genus Radicipes Stearns but so different that it cannot be accommodated within that genus. I now present the following description of this new genus and species in the hope that additional records of it will be forthcoming before the general revision of the family Chrysogorgiidae is published. Family Chrysogorgiidae Verrill, 1883 Distichogorgia, new genus Diagnosis.—Unbranched, flagelliform chrysogorgiids with biserial polyps forming bluntly conical calyces directed obliquely upward. Sclerites exclu- sively flat scales with entire margins and devoid of tuberculate or spinous macrosculpture; distal body sclerites not differentiated as an operculum. Type-species.—Distichogorgia sconsa n. sp., here designated. Comparisons.—This genus differs from Radicipes Stearns in having bi- serial polyps and sclerites exclusively in the form of scales, and from Chal- cogorgia Bayer in lacking specially differentiated opercular scales. The new genus Distichogorgia may be distinguished from the other gen- VOLUME 92, NUMBER 4 877 era at present known in the family Chrysogorgiidae according to the follow- ing key. (Pe GOMeS UNOLANCACE = iis eeeee sci. rh Jlicitatenn drawctamady os We Re 2 SO SIMI SPOT AN CMON 2 aces Ch Alahig Ae iki 's — = S 5 nN 4 < O c } = S aa] ea es = ee 3 DN © Z, al ee ss S © pe Ou VOLUME 92, NUMBER 4 Fig. 3. Distichogorgia sconsa; abaxial and oblique views of single calyx from near apex of holotype (x40). Stereoscopic pairs. blunt, slightly tapered cylinders projecting at roughly 90° (Fig. 1). Between the calyces, the surface of the coenenchyme is somewhat irregular owing to the layer of scales much smaller in size than the supporting scales of the polyps. as Fig. 2. a, Distichogorgia sconsa; calyces (x10) from distal part of holotype, USNM 58444; b, Radicipes pleurocristatus Stearns; calyces (<8) from distal part of colony from Albatross sta. 4969, USNM 49464; c, Radicipes gracilis (Verrill); calyces (<7) from distal part of syntype from Albatross sta. 2037, USNM 8877; d-g, Distichogorgia sconsa; d, supporting scales from calyces (x24); e, smaller interstitial scales from calyces (x 120); f, small scale from tentacle (x 400); g, detail of sculpture of large supporting scale (x 400). 882 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Holotype. —USNM 58444. Length 255 mm excluding holdfast, which was broken off during collection. Diameter of axis just above base, 0.8 mm; about 2 cm from apex, slightly less than 0.2 mm. Color of colony cream white; sclerites colorless, translucent; axis almost white distally, becoming pale straw-colored with moderate metallic iridescence proximally. Holdfast unknown. Comparisons.—Apart from having their polyps in a single row, all known species of Radicipes except R. squamiferus Kukenthal from East Africa have rodlike or fusiform sclerites in the polyps. Examples are the type- species, R. pleurocristatus Stearns (Fig. 2b) from Japan, and R. gracilis (Verrill) (Fig. 2c) from the western Atlantic. R. squamiferus differs from Distichogorgia sconsa in having the scales of the polyps transversely set, not arranged as a bracketlike support, and furnished with conspicuous mar- ginal serrations. Chalcogorgia pellucida Bayer from the Straits of Florida has biserial polyps with smooth, glassy scales, but the sclerites are of con- spicuously different shape and transversely placed, and the polyps are closed apically by distinctly differentiated opercular scales (Bayer, 1949:239, fig. 1; 1956:F216, Fig. 155, la—f). Acknowledgments I am very grateful to Mr. W. R. Brown, chief of the S.E.M. Laboratory of the National Museum of Natural History, Smithsonian Institution, who made the scanning electron micrographs of polyps with a Cambridge Stereo- scan model S4-10 and those of sclerites with a Coates & Welter Cwikscan model 106. Dr. Thomas E. Bowman and Dr. Marian H. Pettibone offered helpful suggestions, for which I extend sincere thanks. This research was supported in part by NSF grant B MS75-07193 to the University of Miami, Frederick M. Bayer, Adjunct Professor, Rosenstiel School of Marine and Atmospheric Science, Principal Investigator. Literature Cited Bayer, Frederick M. 1949. Chalcogorgiinae, a new subfamily of Chrysogorgiidae (Coelenter- ata: Alcyonaria), and a description of Chalcogorgia pellucida, new genus and new species, from the Straits of Florida.—Journal of the Washington Academy of Sciences 39(7):237—240, fig. 1. . 1956. Octocorallia.—/n: Moore, R. C. (Ed.), Treatise on Invertebrate Paleontology. Part F. Coelenterata. Pp. F163-—F 231, figs. 134-162. Geological Society of America and University of Kansas Press. Department of Invertebrate Zoology, National Museum of Natural His- tory, Smithsonian Institution, Washington, D.C. 20560. INFORMATION FOR CONTRIBUTORS Content.—The Proceedings of the Biological Society of Washington contains papers bearing on systematics in the biological sciences (both botany and zoology, including paleontology), and notices of business transacted at meetings of the Society. Except at the direction of the Council, only manuscripts by Society members will be accepted. 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Submit a facsimile with the original and retain an author’s copy. Number pages consecutively at the top. One manuscript page = approximately 2 of a printed page. Underline singly scientific names of genera and lower categories; leave other indications to the editors. Figures and tables, with their legends and headings, should usually be self explanatory, not requiring reference to the text. Indicate their approximate placement by a pencil mark in the margin of the manuscript. Illustrations should be planned in proportions that will efficiently use space on the type bed of the Proceedings (11.8 x 18 cm). Legends require 4 mm of column length per line. Proofs.—Galley proof will be submitted to authors for correction and approval. Changes other than printer’s or editor’s errors may be charged to authors. Reprint orders will be taken with returned proof. CONTENTS a Notes on western Atlantic pipefishes with description of Syngnathus caribbaeus n. sp. and Cosmocampus N. gen. C. E. Dawson On the types in the genus Peloscolex Leidy (Oligochaeta: Tubificidae) Ralph O. Brinkhurst A large parasuchian (phytosaur) from the Upper Triassic portion of the Culpeper Basin of Virginia (USA) Robert E. Weems ~ Comments on the phylogeny of perching birds Alan Feduccia A new genus of water beetle from austral South America (Coleoptera: Hydrophilidae) Paul J. Spangler Neotropical Monogenoidea, 1. Olieapta kruidenieri n. sp. (Axinidae: Indocotylinae) from Thyrinops pachylepis (Gunther) in El Salvador John W. Crane, Delane C. Kritsky, and Robert J. Kayton Bathypontia Sars (Copepoda: Calanoida): eight species, two new, from the Caribbean Sea and Gulf of Mexico Georgiana B. Deevey Description of the larva and pupa of Cylorygmus lineatopunctatus (Coleoptera: Hydro- philidae: Rygmodini) Paul J. Spangler A new genus of madicolous beetles from Ecuador (Coleoptera: Hydrophilidae: Hydro- biinae) Paul J. Spangler A redescription of Scoloplos simplex (Hutchings, 1974) (Polychaeta: Orbiniidae) from Australia Betsy Brown A new genus and species of bopyrid isopod infesting the crangonid shrimp Ponto- philus abyssi Smith in deep water of the northwestern Atlantic Ocean John C. Markham Rectifications in the nomenclature of some Indo-Pacific Littorinidae Winston F. Ponder and Joseph Rosewater Sex-reversal and viviparity in Ophiolepis kieri, n. sp., with notes on viviparous brittle- stars from the Caribbean (Echinodermata: Ophiuroidea) Gordon Hendler Clypeaster kieri, a new species of clypeasteroid (Echinodermata: Echinoidea) from off Bombay, India David L. Pawson and Thomas F. Phelan Hymenaster kieri, a new species of starfish of the family Pterasteridae (Echinoder- mata: Asteroidea) Maureen E. Downey A new crayfish from the Ouachita River basin in Arkansas (Decapoda: Cambaridae) Horton H. Hobbs, Jr. The prostomial pit in Bothrioneurum vejdovskyanum Stole (Oligochaeta): a note on detail revealed by SEM Peter M. Chapman New species of anthurideans from the Cook and Fiji islands (Crustacea: Isopoda: Anthuridea) Brian Kensley The Hydropsyche scalaris group in Virginia, with the description of two new species (Trichoptera: Hydropsychidae) Oliver S. Flint, Jr., J. Reese Voshell, Jr., and Charles R. Parker A redescription of the wood-rasping amphipod Tropichelura gomezi Ortiz, 1976 (Cheluri- dae) from the Florida Keys, with notes on its distribution and ecology James Darwin Thomas The correct name of the helioporan octocoral Lithotelesto micropora Bayer and Muzik | Frederick M. Bayer Distichogorgia sconsa, a new genus and species of chrysogorgiid octocoral (Coelenter- ata: Anthozoa) from the Blake Plateau off northern Florida Frederick M. 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