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WILLIAMS COEDITOR C. W. -HArrT, Jr. ASSOCIATE EDITORS Classical Languages Invertebrates GrorGE C. STEYSKAL THomas E. BowMAN Plants Vertebrates Davin B. LELLINGER LEsLIE W. KNAPP Insects RoBEeERT D. GORDON All correspondence should be addressed to the Biological Society of Washington, Smithsonian Institution Washington, D.C. 20560 ALLEN Press INc. LAWRENCE, KANnsAs 66044 OFFICERS AND COUNCIL of the BIOLOGICAL SOCIETY OF WASHINGTON FOR 1977-1978 OFFICERS President RICHARD S. COWAN Vice President CLYDE F. E. ROPER Secretary W. DUANE HOPE Treasurer OLIVER S. FLINT, JR. Custodian of Publications JOHN H. MILES COUNCIL Elected Members J. LAURENS BARNARD RAYMOND B. MANNING RONALD I. CROMBIE JOHN H. MILES FRANK D. FERRARI iil TABLE OF CONTENTS Volume 90 Abele, Lawrence G. Rediscovery of Sesarma aequatoriale Ortmann, 1894 in the eastern Pacific (Crustacea, Decapoda, Grapsidae) _..._. Anderson, William D., Harry T. Kami, and G. David Johnson. A new genus of Pacific Etelinae (Pisces: Lutjanidae) with redescription of tlre “by pe=Spectes™ ee ee es ee ee ee Baker, James H. Sarsiella pseudospinosa, a new marine ostracod (Myodo- copina; Sarsiellidae) from southern California a Barnard, J. Laurens. A new species of Synchelidium (Crustacea, Am- Phipoda)) trom) sand beaches! in) Californias ee Bayer, Frederick M., and Katherine Margaret Muzik. An Atlantic helio- Poram coral {(Coclenterata) Octocorlha) Bowman, Thomas E. Orchestia vaggala, a new land-hopper from the Galapagos Islands (Crustacea: Amphipoda: Talitridae) Bowman, Thomas E. Ceratolana papuae, a new genus and species of mangrove-boring cirolanid isopod from Papua New Guinea — Brooks, Daniel R., and Robin M. Overstreet. Acanthostome digeneans from the American alligator in the southeastern United States Brown, Walter C., and Angel C. Alcala. A new frog of the genus Rana from! the Philippines! 2.2... MEME 2 ol ee Browning, M. Ralph. The types and type-localities of Oreortyx pictus (Douglas) Vand (Ory. plumiperus” Could! 2 Buden, Donald W., and Darryl L. Felder. Cave shrimps in the Caicos NS | aan) rece ee a) Shad ae ea Cairns, Stephen D. Biological results of the University of Miami deep- sea expeditions. 121. A review of the recent species of Balanophyllia (Anthozoa: Scleractinia) in the western Atlantic, with descriptions of fOUF -MEwW SWECIES: 2a. ee ee Child, C. Allan. On some pycnogonida of French Oceania _........- Child, C. Allan. Four new species of Anoplodactylus (Pycnogonida) from the; westem! North? Atlantiet =: 3... Seat aaeee: : aA ba Coovert, Gary A., and F. Christian Thompson. The Sphegina species of eastern North America’ (Diptera:— Syxphidae)) 2 eee Cowan, Richard S. Studies of tropical American Leguminosae—VIII __ Cressey, Roger F. Two new species of Colobomatus (Copepoda: Cyclo- Podia)iparasiuc. on Lacific fish: ssa fe Crombie, Ronald I. A new species of frog of the genus Eleutherodactylus (Amphibia: Leptodactylidae) from the Cockpit Country of Jamaica __ Dale, George. Apogon mosavi, a new western Atlantic cardinal fish, and a note on the occurrence of Apogon leptocaulus in the Bahamas Deevey, Georgiana B. Conchoecia convexa, a new species of halocyprid ostracod from the Caribbean Sea and Gulf of Mexico _...... Devaney, Dennis M. Ophiomastix koehleri, a new ophiocomid brittlestar (Echinodermata: Ophiuroidea) from the western Indian Ocean ___ Dor, M., and Gerald R. Allen. Neopomacentrus miryae, a new species of pomaceniad fish) fromthe Red: Sea) =a ee ee ee Duellman, William E., and John E. Simmons. A new species of Eleuthero- iD 877-883 975-984 658-668 819-825 1017-1029 669-675 808-8 12 108-115 132-148 440-446 584-596 536-552 237-242 579-583 194-204 19-29 358-366 274-283 183-188 dactylus (Anura: Leptodactylidae) from the Cordillera Oriental of LCISOMM ONG, ASE eA USS PRI a eS Ps ed Darey, RK. .A., and. T. P. @onclaid Postembryonic development and synonymy for Eosentomon rostratum Ewing (Insecta-Protura) — Elbel, Robert E. Buceroemersonia, a new genus of ischnoceran Mallophaga found on the hornbill genus Tockus (Bucerotidae) Emst, Carl H., and James A. Fowler. Taxonomic status of the turtle, Chrysemys picta, in the northern peninsula of Michigan Ferrari, Frank D. A redescription of Oithona dissimilis Lindberg 1940 with a comparison to Oithona hebes Giesbrecht 1891 (Crustacea: Copepoda: Cr clopoiday ee ee ae et A Nt ete Seaton Tt Fisher, Robert D., and Michael A. Bogan. Distributional notes on Notiosorex and Megasorex in western Mexico __......-- Fitzpatrick, J. F., Jr. A new crawfish of the genus Hobbseus from north- east Mississippi, with notes on the origin of the genus (Decapoda, Saimfoantcl ve) meme ee TN Reet iin ere ade Ten on Te oD ah ee Flint, Oliver S., Jr., and Joaquin Bueno-Soria. Studies of Neotropical cad- disflies, XXI. The genus Lepidostoma (Trichoptera: Lepidostomatidae ) Gloyd, Howard K. Descriptions of new taxa of crotalid snakes from Olnmamancm@eylone( Sri Tuanka))| ces ee hg Gordon, Robert D. ... AO NA ee EEE eee Ceratolana _______ papuae _- Colobomatus* springen). 2s oe ee ee ee EEE eee colletei ___ @olomastix janiéeae: 2.2 ee EE Conchoecia conve NL se oe eee GO TEC LAUT ee led a la ah Ss penascoens DS ih a a a tg hy at tana I Cycloleberis. christiGi 2.22: oe eller Dactylocythére-astraphes 22 eee COTDUS ae promincula spinescens Distosquilla Faxonella blairi _ Gammarella, fucicola n; ¢.. . e eeeeee Hobbseus petilus Lenisquilla Letisquilla Orchestia vaggala SeEMBRNIGMES ILL OC CIO (I Samanta eet St Ree te ce eT oe ee anand lk cently, 2 Parthenope (Platylambrus) granulata n. c. TYSONS GCM aes ee aN ee eee er OO ee ae Th ers PDQ. ek Sete SS Ea URIS CE ESS Da Oe eee Sreeioeplnen® OCU GT Tea i 0 NaI el ch te Ue LT Procambarus (Scapulicambarus ) strenthi grein (Qing: see Nee SS ES I ei eee cet oe ae ee (DEO DS DOOR. ets os ARN es Py ae eS Tie a el ene eee PN DEDW DSO, es ps USI eR eg ol eG ac GEMMA TOLTN “10, sO, Ce eae I eae Ee An Noes dO ne Als Lys vn \ pe Oe Reams £ ON ee ree Ral Ba ORTOOITOCES 1 CX Mma tay sea et ee ee TTS ATG CT Teale oy (Oe seek to an lea SAA Eo a Le a I alee a eee Pemteelne li GiviIMENECTOMUCOM - esa asd ee Baas 171 1) 5 EL Sn Ia he nat Se Mt us COPDULIS ss EES ee Re eR OEE ac eR Nc Re OR eo Spmialle, BgOFH a cS Oe OO DIEU, ace cess ta gt le ee Se DIR GILONDG, cp ga ce A @es SOGUSUVGHER “inspec e e LEME DUDIE ase ag Oa na LEAS Saal. YICTODSSOD. aia gat ee ln ae Re LT OnE eT a Scenics aba aanyas ee RS aE eee eR ee Ea aCOCI ICTS OI Ome Nee eee Me ee OAS RA Or TO eek pee isl ATC. 11 ne tine eg he ree oe ee ee ty GY, RS oe See ee UGA OD aerate cE ROSEN a 2 Ne (DLA eed COC Ut Py IO NRE SMe eg a aa EE es ale SC ce el ae CU Lg ee WAR Le svn Ey el ig a A SINS Oe We OL a Ane NS DN ie enetgkes Broepais 22S ee rare ee ee wees Eon OSKOMMAMCEZEC CULT U me aeeN saea A ee ee ee Pee he ls eo GONE cece DI ar Pe nN ee eS aan ROLUOSOPG: meas ee sh A NED Di a OO ORE atc Te Tn eae I ae SCT AUATLOGGE | LSS SIDE TES TIS EAR SU lt Ne nS an PY Ps ECLCETTLCLING C10 SC Maen a ean cE eS PUD erties se te oe ee BETA MINMNONAMATACOCCTAS -. wee ee UBTOPD OS. etcaccce She AAR act ee SER ea Oe ee era eRe _Sphegina (Sphegina) appalachiensis ECHINODERMATA Ophiuroidea BMGT IN ACLU 79 CECE ce Mminomastix Koehler ___........_....____-.-...... CHORDATA BROT METRO Si eee 20 Eyphessobry.con. .diancysirus, 0.6. ee eee 349 SOCOOfi 8 A ee Sin 327 Kidausewitzia...aphanes.. 2. 2 eee ssl INeopomacentrusiminjad. ee eee 183 Rercinasantesela,.2 3 eer 7 Randallichthys:....2 5. le ee 1 id oe 89 FU ATMEMETOSUS GMC i a 91 Amphibia Eleutherodactylus. chiastonotus ...... Eee 425 UNC <2. Son a ee ew A Oe 60 SIS PROGCMUS - eEeeeeeeeeeeeeeeee 194 ZOuctOtylus ee eee 432 RRO SOR Gta eo 669 Reptilia Agkistrodon blombhoffi siniticus 0... eee 1004 blomhothi dubitatus, 222. whee Eee 1007 halys’ Coonatus: = 22 n r 1002 iyjomale: quali. 2 he eo ee 1011 Sphacrodactylus ‘micropithecus 2. ET—E—E Eee 986 TANG TGs 2 ee 247 TANG MCEIOTIUS! 6s be ee 248 TANG STONE 2 ee Dell Mammalia Grocidura horstieldi kurodae ee 461 Tussula Noslethi 2 lee 465 Microtus ochrogaster similis... eee 49 Ringuimis alfrednewtoni 2 eee 690 NCTICUD IS eee ee eT ee a ee 448 XUMPUheCus, Lo 0 Ne ee eee 449 XID m7, 06/5 Proceedings of the BIOLOGICAL SOCIETY of WASHINGTON Volume 90 16 June 1977 Number THE BIOLOGICAL SOCIETY OF WASHINGTON 1977-1978 Officers President: Richard S. Cowan Secretary: W. Duane Hope Vice President: Clyde F. E. Roper Treasurer: Oliver S. Flint, Jr. Custodian of Publications: John H. Miles Elected Council J. Laurens Bamard Raymond B. Manning Ronald I. Crombie John H. Miles Frank D. Ferrari PROCEEDINGS Editor: Austin B. Williams - Coeditor: C. W. Hart, Jr. Associate Editors Classical Languages: George C. Steyskal Invertebrates: Thomas E. Bowman Plants: David B. Lellinger Vertebrates: Leslie W. Knapp 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. Cor- respondence concerning membership should be addressed to the Treasurer, Biological Society of Washington, National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560. The Proceedings is issued quarterly in February, May, August and November. Manuscripts, corrected proofs, editorial questions should be sent to the Editor, Biologi- cal Society of Washington, National Museum of Natural History, Smithsonian Institution, Washington, 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 Application to mail at second class postage rates is pending at Washington, D.C., and additional mailing office. EDITORIAL Inflation continues to increase printing costs. To meet costs for the Proceedings, the Biological Society has already raised annual dues and voluntary page charges but finds that these increases will not be enough to sustain publication at the present standards of quality and number of pages per volume. Immediate raises can be avoided and future ones perhaps held to a minimum by accepting an Allen Press proposal that we adopt, with several small journals, a standard format for mass production printing. The Council has voted this acceptance, knowing that some changes in the familiar journal would result. Since its inception in 1882, the Proceedings has had an unchanged page size that is most economical for short-run journals, virtually the same type bed, and only minor variations in type, usually 10 point for introductory material, 8 point for species accounts, legends, tables and literature cited, and 6 point for footnotes. The new format retains the same page size but increases the line length from 23 to 28 picas, and the page length from 39 to 43 picas by changing the running heads. More space is thus provided for both text and illustrations. Composition is simplified by setting all text and abstracts in 10-point type, and figure legends, tables and literature cited in 8-point type. Abstracts, never accepted previously, are now considered in- creasingly useful adjuncts of even short descriptive papers because they condense information for the abstracting services. Boldface type, un- fortunately, is no longer available for announcing new taxa. Rather than individually saddle stapled articles being issued irregularly in occasional paper style, articles will now be issued four times per year as a bound journal. Paper is changed from the former 70-lb enamel or matte finish to 60-lb enamel finish. Author's addresses are placed at the end of articles. These changes represent some losses, but there are more gains. Authors will benefit from the enlarged type bed and fewer pages per paper. Editing, all done by volunteers, is simplified. Articles in a bound journal do not have to be sorted for mailing by the Custodian of Documents, also a volunteer, nor do the reprints have to be mailed by him. Bound journals issued four times a year can be mailed second class. Information tor contributors and readers can be placed on the cover. It is our hope that the newly designed journal is pleasing, more useful, and in the long run more economical than the style that served well for so long. The journal cover offers space for a distinctive logo, one that describes the scope of the Proceedings and/or the situs for the Society. We invite suggestions for an appropriate symbol to be selected by the Council and rendered by a designer. Austin B. Williams Editor PROC. BIOL. SOC. WASH. 90(1), pp. 1-5 FAXONELLA BLAIRI, A NEW CRAWFISH FROM THE RED RIVER DRAINAGE OF OKLAHOMA AND ARKANSAS William A. Hayes and Rollin D. Reimer The genus Faxonella (Creaser, 1933) represents a small assemblage of crawlishes ranging in the Gulf coastal plain from South Carolina and Georgia through Florida, Alabama, Mississippi, Louisiana, Missouri, Ar- kansas, Oklahoma, and Texas (Creaser, 1933; Fitzpatrick, 1963; Hobbs, 1942, 1969). This is the fourth species to be assigned to Faxonella. The material on which it is based came from the Little River drainage of southeastern Oklahoma and the Red River drainage below the confluence of these two streams in southeastern Arkansas. The southern boundary of this species in the Red River drainage has not yet been determined. Faxonella blairi, new species Figs. 1-7 Faxonella clypeata.—Fitzpatrick, 1963:61 [in part]. Diagnosis.—Lateral rostral spines and cervical spines absent; rostrum short, broad (less than 1.5 times as long as broad). Areola about 3 times as long as wide (length 30% of carapace length). Male with hooks on third pereiopods only. First pleopod of Form I male extending cephalad to coxop- odite of first pereiopod, terminating in 2 rami; both rami twisted mesially, overlapping at midline of sternum. Central projection long (about 6 times longer than mesial process), slender, bladelike, distal half recurved mesially with tip bent caudally, corneous; mesial process short, slender, cylindrical, recurved caudally, noncorneous. Overlapping of terminal elements in normal position about % distance from base of central projections, at level of tips of mesial processes. Annulus ventralis immovable, subquadrate, with 2 elevations on cephalic half and prominent subangular caudal lip. Holotypic male (Form I).—Body subovate, compressed laterally. Width of thorax greater than that of abdomen (7.0-6.2 mm respectively). Height of carapace greater than width (7.3-7.0 mm respectively). Greatest width of carapace slightly posterior to midway between caudodorsal margin of cervical groove and caudodorsal margin of carapace. Areola moderately wide, 3-5 times as long as wide. Length of areola 30% entire length of carapace. Rostrum reaching end of second segment of antennule; slightly concave, depressed with convergent sides; low lateral ridge without lateral spines or tubercles. Acumen subtriangular. Postorbital ridges merging PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Tf Figs. 1-7. Faxonella blairi, n. sp.: 1, Carapace of holotype, dorsal; 2, Annulus ventralis of allotype; 3, Epistome of holotype; 4, Chela and carpus of holotype, dorsal; 5, Carapace of holotype, lateral; 6, Right antennal scale of holotype; 7, Gonopods (right, except c): a, e, Form II ¢; b, c, d, Holotype; a, b, Lateral; c, Ventral; d, e, Mesial. VOLUME 90, NUMBER 1 3 cephalically into carapace. Branchiostegal spine present. Cervical spines absent. Length of abdomen equal to that of thorax. Anterior section of telson with 2 spines in each posterolateral corner. Antennae slightly longer than cephalothorax. Antennal scale extending to midlength of third segment of antennular peduncle; broadest at mid- length; weak spine on outer distal margin. Palm of chela of first pereiopod inflated, subcylindrical, with ridge of tubercles on mesial surface; dorsal surface covered with smaller tubercles. Movable finger slightly longer than mesial surface of palm. Fingers flattened, each with 4 rows of incisor-like tubercles on opposing surfaces, sparsely setose along inner surfaces. Gape between fingers absent. Carpus longer than broad, tuberculate along mesial margin, shallow longitudinal furrow above. Merus with 2 reduced spines on upper distal surface; lower surface with mesial row of 12 tubercles; lateral irregular row of 7 tubercles, row of 5 reduced tubercles on distal border of lower surface. Ischiopodite of third pereiopod with prominent hook, latter corneous distally. First pleopod extending cephalically to coxopodite of first pereiopod when abdomen flexed and terminating in 2 distinct processes, tips of both curving slightly caudally. Central projection long, slender, blade-like, twisted mesially in proximal half with distal half pointing cephalolaterally (in normal position), and corneous; mesial process short, slender, also bent cephalomesially, and noncorneous. Terminal elements overlapping, in normal position, about % distance from proximal end of central projection, at level of tips of mesial processes. Female allotype—tThe allotype differs from the holotype in that the chelae are shorter in relation to carapace length and flattened dorso- ventrally to a greater degree than in the holotype. Annulus ventralis subquadrate in outline with extended caudal lip; firmly embedded in the sternum; cephalic portion partly covered by two elevations on cephalic half. Surface contours and sinus as illustrated (Fig. 2), not so complex as in F. creaseri or F. clypeata. Type-locality—Woodland swamp approximately 100 yards W of U.S. Hwy. 259 on N side of the Little River, 7 mi S of Broken Box, McCurtain County, Oklahoma. Individuals were common in swampy areas and in temporary pools in roadside ditches. The water in the area was static, turbid, shaded, and had an average depth of 12 inches. Crawfish associates collected at the type-locality were Procambarus acutus acutus (Girard, 1852), and Fallicambarus hedgpethi (Hobbs, 1948). Disposition of types—The holotypic male and allotypic female are deposited in the National Museum of Natural History, Smithsonian Insti- tution (USNM 145705 and 145706, respectively ). 4 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON a eS eee Measurements (in mm ) Holotype Allotype i ot ee lee Carapace Height Thess) 9.1 Width 7.0 8.9 Length 15.7 19.8 Areola Length 4.7 6.2 Width 0.8 LF Rostrum Length 3.0 4.9 Width 2.9 3.0 Chela Length of inner margin of palm 4.8 4.0 Width of palm 3.0 3.8 Length of outer margin of hand 10.2 9.4 Length of dactyl 4.4 49 Antenna Length 22.0 19.4 Antennal scale Length 3.2 S15) Gonopod Total length 7.0 Central projection (Length) Al Mesial process (Length) 0.8 Annulus ventralis Length 1.9 Width 2.5 Range.—We have examined specimens of F. blairi from three localities in southeastern Oklahoma and southwestern Arkansas other than the type- locality: OKLAHOMA. McCurtain Co.: 4 mi N of America, near Shine- well (roadside slough); Forked Lake, SE of Broken Bow. ARKANSAS. Sevier Co.: West Otis (large pond near Oklahoma state line). Relationships—Faxonella blairi appears to be a geographic isolate of the genus in the Red River drainage of southeastern Oklahoma and south- western Arkansas. The annulus ventralis of F. blairi is firmly embedded in the sternum, much more than in other species of Faxonella, and the sinus simpler in sculpture. In these features, this crayfish is more similar to Faxonella creaseri than to either F. clypeata or F. beyeri. The females of F. blairi can easily be distinguished from other species in the genus Faxonella by the prominent caudal lip of the annulus ventralis. The males can be distinguished from all other Faxonella species by the more nearly straight central projection of F. blairi which reaches to the coxae of the first pereiopod. This character also points to a close relation- ship with F. creaseri in which the central projection reaches basically to the same level but the distal half of the ramus is bent more mesially. In Faxonella clypeata and F. beyeri, the mesial process is much shorter and more curved. VOLUME 90, NUMBER 1 5 Etymology.—We are pleased to name this crawfish in honor of Dr. Albert Patrick Blair, Department of Life Sciences, University of Tulsa, Tulsa, Okla- homa. Dr. Blair has contributed many collections of crawfishes from the southwest to various museums and thus has greatly increased our knowledge of these animals. Acknowledgments The authors thank Dr. Horton H. Hobbs, Jr., National Museum of Natural History, Smithsonian Institution, for many courtesies during the preparation of this description. Literature Cited Creaser, FE. P. 1933. Descriptions of some new and poorly known species of North American crayfishes. Occ. Pap. Mus. Zool. Univ. Michigan 275:1-21. Fitzpatrick, J. F., Jr. 1963. Geographic variation in the crawfish Faxonella clypeata (Hay) with the definition and defense of the genus Faxonella Creaser (Decapoda, Astacidae). Tulane Stud. Zool. 10:57-79. Girard, C. 1852. A revision of the North American Astaci, with observations on their habits and geographical distribution. Proc. Acad. Nat. Sci., Philadelphia 6:87-91. Hobbs, H. H., Jr. 1942. The crayfishes of Florida. Univ. Florida Pub., Biol. Series 3:1-179. —. 1948. A new crayfish of the genus Cambarus from Texas, with notes on the distribution of Cambarus fodiens (Cottle). Proc. U.S. Nat. Mus. 98:223-231. 1969. On the distribution and phylogeny of the crayfish genus Cambarus. Rpy 93-178 in P. GC. Holt, R: L. Hoffman, and C. W. Hart; Jr, eds. The dis- tributional history of the Biota of the Souther Appalachians, Part I: Inverte- brates. Research Division Monograph, Virginia Polytechnic Institute, Blacksburg, Virginia. Department of Wildlife and Fisheries Sciences, Texas A&M University, College Station, TX 77843. Present address —(WAH_) Department of Science, Murray State College. Tishomingo, OK 73460. PROC. BIOL. SOC. WASH. 90(1), pp. 6-18 PERCINA (IMOSTOMA) ANTESELLA, A NEW PERCID FISH FROM THE COOSA RIVER SYSTEM IN TENNESSEE AND GEORGIA James D. Williams and David A. Etnier Introduction Like Percina lenticula Richards and Knapp, Percina antesella was first taken in a tributary to the Etowah River (Coosa Drainage), Cherokee County, Georgia, in an area that is now flooded by an Army Corps of Engi- neers impoundment, Alatoona Reservoir. The collection, of three specimens, was made by Donald C. Scott and O. Tyson in Shoal Creek, 5 July 1948. One specimen was sent to the Museum of Zoology, University of Michigan; the remaining two were deposited in the University of Georgia fish collec- tion. While visiting the University of Georgia collection in 1966, the two specimens were examined and recognized as a new species. Attempts by Williams to secure additional specimens from tributaries to Alatoona Reser- voir in Cherokee County, Georgia, were unsuccessful. In 1968 David Etnier and Robert Stiles of the University of Tennessee collected additional specimens in the Conasauga River (Coosa Drainage), Polk and Bradley counties, Tennessee. Since 1968 approximately 70 speci- mens have been collected from three localities on the Conasauga River in Murray County, Georgia and Polk and Bradley counties, Tennessee. The three localities are within a 15 mile section of the Conasauga River. The new species, Percina antesella, brings to five the number of species belong to the subgenus Imostoma. Four of the species (P. antesella, P. ouachitae, P. tanasi, and P. uranidea) appear to represent a distinct phyletic line, characterized, in part, by the presence of four or five dark saddles across the dorsum. The fifth species of the subgenus Imostoma, Percina shumardi, lacks saddles. Within the “saddleback” species group it appears that Percina tanasi and Percina uranidea form a species pair, while P. ouachitae is interpreted as a more recently evolved generalized species capable of adapting to a variety of stream conditions. Percina antesella is considered a relict species which is not closely aligned with any of the other three species. The recognition of Percina ouachitae (Jordan and Gilbert) as a distinct species rather than a synonym of P. uranidea (Jordan and Gilbert) is based on unpublished studies by Bruce A. Thompson and Robert C. Cashner. Their validation of P. ouachitae and a discussion of relationships within the sub- genus Imostoma will appear elsewhere. We acknowledge with thanks the assistance of the following individuals. For the collection and loan of material we are grateful to R. M. Bailey, — SR VOLUME 90, NUMBER 1 7 Museum of Zoology, University of Michigan (UMMZ); H. T. Boschung, T. S. Jandebeur, and H. Harima, University of Alabama Ichthyological Col- lection (UAIC); G. H. Clemmer, Mississippi State University (MSU); N. H. Douglas, Northeast Louisiana University (NLU); E. A. Lachner, National Museum of Natural History (USNM); J. S. Ramsey, Auburn University (AU); M. E. Sisk, Murray State University Collection (MSUC); D. C. Scott, University of Georgia (UG); W. M. Howell and R. A. Stiles, Samford University; W. C. Starnes, University of Tennessee (UT); R. D. Suttkus, Tulane University (TU); and R. W. Yerger, Florida State University (FSU ). We especially thank Robert C. Cashner and Bruce A. Thompson for their valuable comments on the manuscript during its preparation. The methods of Hubbs and Lagler (1958) were followed in obtaining counts and measurements. Vertebral counts followed the methods of Bailey and Gosline (1955). Percina antesella, new species Amber Darter Iie, I Percina uranidea. Richards and Knapp, 1964:700 ( misidentification, supple- mentary material, UMMZ 156697). Percina (Imostoma) sp. Stiles and Etnier, 1971:12, 15 (Conasauga River, Polk and Bradley counties, Tennessee); Page and Whitt, 1973a:3, 4 (Conasauga River, Bradley Co., Tennessee); Page and Whitt, 1973b:614- 616, 621 (Conasauga River, Bradley Co., Tennessee); Page, 1974:66-72, 75-82, 85 (Conasauga River, Bradley Co., Tennessee ). Percina sp. of uranidea. Dahlberg and Scott, 1971:47, 64 (Coosa System, Georgia and Tennessee). Percina sp. Narrowsaddle Darter. Miller, 1972:245, 246, 250 (Rare, Cona- sauga River, Georgia and Tennessee ). Percina sp. Etnier, 1975:472, 482, 484 (key to species of subgenus Imostoma and comparison with P. tanasi). Percina (Imostoma) sp. Page, 1976:258, 261 (Upper Alabama River Sys- tem). Holotype —Adult male, TU 94031, 47.0 mm standard length (SL), Cona- sauga River, Tennessee Hwy. 74, Bradley Co., Tennessee 9-10 April 1970. Paratopotypes.—UT 91.471 (10 specimens), collected with the holotype, TU 58938 (7), 17 Oct. 1969; TU 58968 (3), 19 Oct. 1969; TU 65937 (7), 299 June 1970; TU 69143 (8), 15 April 1971; TU 78368 (1), 1 July 1972; UAIC 4459.01 (1), 6 March 1971; UAIC 4729.01 (11), 29 Nov. 1969; USNM 210666 (1), 16 May 1970; USNM 210667 (3), 22 Oct. 1969; USNM 910668 (2), 22 Feb. 1970; UT 91.799 (2), 28 June 1973; UT 91.800 (1), 22 July 1972; UT 91.1100 (1), 4 Nov. 1973. 8 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 1. Percina antesella n. sp., paratopotypes. Top: UT 91.799, ¢, 53 mm SL. Middle: UAIC 4729.01, 2, 52 mm SL. Bottom: UT 91.1100, ¢, 52 mm SL. Other paratypes.—Georgia, Cherokee Co.: UMMZ 156697 (1) and UMMZ 194317 (2), Shoal Creek, first bridge above mouth (now flooded by Ala- toona Reservoir), 5 July 1948. Murray Co.: TU 59427 (7) Conasauga River, Hwy. 286 bridge, 2.3 mi W of junction of Hwy. 225 and 286, 19 Oct. 1969. Tennessee, Polk Co.: UAIC 4730.01 (1) Conasauga River at Boanerges Church bridge, 3-4 Nov. 1968; USNM 210669 (1) Conasauga River, second riffle below Boanerges Church bridge (first bridge above Tennessee Hwy. 74 bridge), 11 Oct. 1969. Diagnosis.—A species of the genus Percina, subgenus Imostoma (see Page, 1974:85), distinguished from known species by: anterior saddle on pos- terior portion of nape, entirely anterior to spinous dorsal fin (under anterior portion of spinous dorsal fin in Percina tanasi, P. ouachitae, and P. uranidea); lower procurrent caudal rays of breeding males extended ventrally pro- ducing a triangular flange bearing breeding tubercles (no such flange in ! | 9 VOLUME 90, NUMBER I LP'9 C1G-061 SL'00G 99ST FEC-PV8T B8LE0G 168 O€S-E6I GLOIG 5 OG epeccinle 0S FO E901 GSS-E6I Y9E'80Z OOIL S8E2-S6I SZGIZ ? Yue, UlY Aad SIFI SES-Z8I §6'0% SII PIG-9LI SP'86I GOI LES-O8T LO'TIIZ é (ysu9| SAL ASNT Seal POTL LIZ-O8I 00861 6S°9 iZGe lO CEG SOIC P apounped [epne) og’ €8-89 Cr'eL Lg PS-IL LOS. 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I¥'88 é LEE oa!) 06'S COT 06-91 1618 gee S6-18 G)'L8 2 Yysue, yous IF’ 962-686 SS'PLZ 09°8 G6Z-08Z FE F8Z aa FOE-SLZ -GO'SES é 918 OO€-6SG SF'SLZ Wile LOS=C9G. 1GISIG LYS OOk-F8Z 0'S6Z P Ysue] peo] £9'C O19-OZF_ FE LF L169 OLS-9EE LE'OF OL'S Z6S-OSE 90'°9F j cT'9 SF9-8ZF «O86 C6'F GOG-GLE = LO' PF 91'S G8St-FIF 60°SF p Yysue] plepuris (606: 20Z) (86:2 TIT) (BCE SOL) JSBOD FIND apzyoono *g ‘YI eCSSTP, 29 “UAT, apyyooNo * J "ds ‘U D)/aS9}UD * J | ‘UOTeIAVp pilepurjys = 7S ‘sonyeA Fo ssuel = (A ‘uve = X *(}aq[Ig) pue uepIO[) aDpYODNO ‘g pue sSaloads MoU DIaSajUD DUIOIag JO "TS JO SpuBsnoYy} Ul S}JUSWoINseaIY “T 9[qeL PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 10 PE TI VS-9P CT'6S vO'6 89-9F 8L°S¢ g9°9 69-IF 9T'1TS re) 9[ppes GEOL CL-8E CPV 9c $9'6 C9-8E 9E°9¢G £68 £9-66 80°87 ’ puooss jO YAPIM COSI 6IP-8SPE §=OL OLE GV LI C8t-ITe& 9G 19E LOVT OLE-LOE Y99'8EE } a[ppes LL OT FORP-GEE = =0G'69E LOST c8t-9EE OO9CE I9 OL 6PE-9TE G6 PES ’ puoses 0} yndID90Q 09'GT S6-PP 06°89 OLS 98-19 SL°GL £9°9 9S-8G OS PP } 9[PPpes Sait €6-TS O09 TL 866 O8-LS 97°99 egg 0S-9€ OG EP £ SUF JO YIPIM PV GL 69I-LGI c9'9OFT O6'L POT-OFI SLES 86'S 88-69 I8'GL } Q[PPes SUF 09ST 99T-IGI OS PFI SUGI POT-OFT LO VST cc’9 O8—-LS CO GL ’ 07 ynd1D99 G8 PV VL-VS cL'S9 cV'9 G8-L9 OF PL 699 F6-£9 GG E8 6 ourds yeue GIL 88-€9 cO'TL VV 9 P8-€9 PIEL 86'S TOI-62 0S'06 2 jo Yue IOGL I8G-8EG 06096 CV's LLG-6FG LL 696 LOSPF 966-CGSG FE ILIG } Yyysues] uly Cer O€t-OLG OV ETE VG OL €6E-98G LOPOE LOSE ITE-GG6 = GV LLG 7 [estop 30S 69 GI 16G-6FSG G9 OLD GV GI C8c-EFG 00° LGG OLOT 69G-OEG POPVVG fo) yysus] Wy CcLel OO&S-9FG OF SLE 09'0T G8G-GVG 9IE'8SG 866 CS9IG-LGG C8 PPG e [esiop Aurdg re M x OS M x S M » *penut}uor) VOLUME 90, NUMBER 1 1 Percina tanasi, P. ouachitae and P. uranidae ); transverse scale rows 15-19, usually 16 or 17 (11-16, usually 13-15 in P. ouachitae ). Description.—Percina antesella is a small, moderately slender species rarely exceeding 60 mm standard length (SL). Head length contained 3.4- 3.6 times in standard length. Head length in P. antesella slightly longer than in P. ouachitae, due in part to the somewhat longer snout of P. ante- sella compared to that of P. ouachitae. Snout moderately long and pointed, slightly longer than orbit. As in most species of subgenus Imostoma, eyes placed high on head and interorbital width very narrow. The general body shape is illustrated in Fig. 1. Proportional measurements of 44 paratypes are presented in Table 1. Gill membranes usually separate, narrowly connected in some individuals. Development of frenum variable but usually incomplete. Cephalic sensory canal system complete with three pores in supratemporal canal, four pores in supraorbital, and a single coronal pore. Preoperculomandibular canal usually has 10 pores. Infraorbital canal has 8 or 9 pores, typically 8. Dorsal spines range from 9-11 with mode of 10; dorsal rays range from 12-15, usually 13 or 14. Pectoral rays range from 13-15, usually 14. Anal rays usually 9 or 10. Typically 12 branchiostegal rays, six per side. Vertebral counts varied from 38-40 with mode of 39. Frequency distributions of fin- ray and vertebral counts are given in Table 2. Body squamation usually complete, except anterior portion of nape and belly, both of which vary from naked to almost complete coverage with em- bedded scales. In males, midventral row of modified scales not well de- veloped, width usually only slightly larger than scales of adjacent rows. Males usually have two modified scales present between pelvic fins and an additional three to seven scales along ventral midline of posterior half of breast. A few scattered embedded cycloid scales present on breast of some individuals. Opercles usually only partially scaled, and cheeks typically naked. Lateral line complete with 51-66 scales, usually 55-61. Usually 16- 18 transverse scale rows and 19-21 caudal peduncle scale rows. Frequency distributions of scale counts are given in Table 3. Coloration in life —On trunk, golden brown of dorsum extends ventrad to first or second scale row above lateral line. Four dark brown saddles present on dorsum, first crosses posterior part of nape, entirely anterior to origin of spinous dorsal fin. Posterior margin of this saddle usually one to three scales anterior to first dorsal spine and saddle usually four to six scales in width. In Percina ouachitae first dorsal saddle beneath anterior portion of spinous dorsal fin (Fig. 2). Second dorsal saddle situated under pos- terior end of first dorsal fin. Second saddle usually with anterior margin between last two dorsal spines and posterior margin at or just anterior to soft dorsal origin. Third dorsal saddle usually under three or four rays between the sixth and eleventh dorsal rays. Fourth dorsal saddle on pos 12 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Table 2. Frequency distributions of fin-ray and vertebrate counts in Percina antesella and P. ouachitae from drainages east of the Mississippi River. Dorsal spines 9 10 11 12 x P. antesella 2 43 25 10.33 P. ouachitae i) Are) yA! 6 10.35 Dorsal rays 12 13 14 15S x P. antesella 9) 38 22 1 eo) P. ouachitae 8 109 283 83 13.91 Pectoral rays 1S} 14 15) 16 17 x P. antesella 19 47 4 13.79 P. ouachitae Se OA Gi al i 14.60 Anal rays 8 9 10 WIL 12 x P. antesella 1 32 31 9.34 P. ouachitae 18 VANS) 257 26 10.31 Principal caudal rays 14 155 16 17 x P. antesella 1153 44 7 15.88 P. ouachitae 2 Cie b4 27 15 557/ Vertebrae 38 39 40 x P. antesella 8 37 3 38.90 P. ouachitae 10 45 7 38.95 terior portion of caudal peduncle, just anterior to procurrent caudal rays; usually three to four scales wide at dorsal midline. All four dorsal saddles angle anteriad, narrowest at dorsal midline, expanded ventrad, terminating near lateral line. Ventral expansion of saddles most prominent on second and third saddle. Pigment along lateral line present in varying concentrations. On anterior third to half of body lateral pigment consists of irregular light brown blotches scattered just below lateral-line scale row. Light brown to brown irregularly shaped blotches present on or just below lateral-line scale row on posterior half to two-thirds of body. Most prominent blotches are on VOLUME 90, NUMBER 1 13 Table 3. Frequency distributions of scale counts in Percina antesella and Percina ouachitae from drainages east of the Mississippi River. Lateral line scales 46 47 48 49 50 51 52 53 54 55 56 57 58 59 60 61 62 63 64 65 66 x P. antesella il Se) EGP Miao Ou cGl ly comin Seek 56.65 P. ouachitae Qe OeLONI 3835246) 05) 6016959 50lo>) 20,10) oat SP a Transverse scale rows Ike IDA IS ar sy IS alee ALS} JI) X P. antesella 4 OD HO)» Bl 16.56 P. ouachitae 2, 4 102 201 96 10 14.06 Caudal peduncle scales We Ike IQ) BO GAN oe Xx P. antesella Do IS BO SX) Tl 20.04 P. ouachitae 24 83 191 110 28 2 19.11 posterior part of body and usually appear as irregular expansions along ventral margin of dorsal saddles. Below lateral pigment, concentrations of dark chromatophores present on groups of one to three scales on ventro- lateral body surfaces, most apparent on posterior third to half of body. Venter pale yellow to cream color. Dorsal and dorsolateral surfaces of head, including upper portion of cheeks, opercles, and snout tip dark olive brown. Lateral and ventral areas of head yellowish to cream color. Interopercle and subopercle iridescent pale blue. A prominent dark brown subocular bar about width of pupil slopes downward and slightly forward from eye. A small concentration of melanophores usually present on gular region at junction of branchiostegal membranes. Spinous dorsal has vague gray black basal and marginal bands with central third to half of fin clear. Chromatophores apparently absent from spinous dorsal fin. Soft dorsal fin has four or five clusters of dark chromatophores evenly spaced along each ray, usually best developed toward base of fin; membranes unpigmented. Caudal and pectoral fins similar to soft dorsal but usually have a somewhat irregular pattern which gives them a more mottled appearance. A caudal spot of varying intensities usually present near middle of caudal fin base. Unpigmented areas usually present above and below caudal spot. Anal and pelvic fins unpigmented except for an oc- casional cluster of dark chromatophores. Breeding tubercles—Males of Percina antesella taken in April have well developed tubercles which are different in their distribution and somewhat 14 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 2. Percina ouachitae (Jordan & Gilbert), UT 91.860, Buffalo River approxi- mately 1.3 mi above its mouth, Humphreys Co., Tennessee, 10 November 1973. Top: 6, 44 mm SL. Middle: 2,53 mm SL. Bottom: 92, 50 mm SL. reduced in size compared to P. ouachitae and P. uranidea. Tubercles are present on the caudal, anal, pelvic, and pectoral fins and some scales on venter, breast, and caudal peduncle. Percina antesella lacks tubercles on branchiostegals, rami of lower jaw and scales on cheeks and opercles in contrast to P. ouachitae which has tubercles on these areas. In males, caudal fin has well developed tubercles on unbranched and branched portions of ventral fifth to eighth rays. Posterior third to fifth lower procurrent caudal rays extended ventrally into a triangular flange which may have up to four tubercles along base. Both P. ouachitae and P. uranidea lack the triangular flange. Three to six tubercles present on both spines of anal fin. Tubercles present on distal half to two-thirds of all anal rays. Pelvic fins have tubercles on dorsal and ventral surfaces of rays, usually best developed on ventral surface of rays, from near base to fin margin. Distal half to two-thirds of ventral surface of pelvic spine usually with a ridgelike structure; occasionally two or three low, rounded tubercles present on proximal third of spine; tubercles usually absent from dorsal surface of spine. Ventral third of pectoral fins with a few small tubercles on distal half of longer rays. VOLUME 90, NUMBER 1 15 ee S pt. 85 Cau if 37° ‘Simpsonm@on aU 43984 (1). TCHEFUNCTE RIVER SYSTEM. Louisiana: Washington Parish: NLU 6455 (1). AMITE RIVER SYSTEM. Louisiana: Livingston Parish: TU 29047 (1). East Feliciana Parish: TU 75550 (8). Mississippi: Amite Co.: NLU 11601 (4). Mississippi River tributaries, Mississippi—THOMPSON CREEK SYS- TEM. Louisiana: West Feliciana Parish: TU 55303 (1), TU 69616 (24). Mississippi: Wilkinson Co.: TU 74706 (53), TU 74774 (30). BAYOU SARA SYSTEM. Mississippi: Wilkinson Co.: TU 59998 (1). HOMOCHITTO RIVER SYSTEM. Mississippi: Lincoln Co.: TU 71206 (1), TU 73271 (1), TU 78054 (1). COLES CREEK SYSTEM. Mississippi: Jefferson Co.: TU 50456 (1). BAYOU PIERRE SYSTEM. Mississippi: Claiborne Co.: FSU 9221 (2). Copiah Co.: FSU 9274 (3), TU 76810 (2), TU 80270 (31), USNM 266609 (6). Mississippi River Tributaries, Tennessee and Kentucky—HATCHIE RIVER SYSTEM. Tennessee: Hardeman Co.: USNM 190786 (1). Mc- Nairy Co:: USNM 190755 (2), UT 91.798 (8). Tipton Cols Use 9N60r (24). MISSISSIPPI RIVER PROPER. Tennessee: Tipton Co.: UT 91.563 (1). OBION CREEK SYSTEM. Kentucky: Carlisle Co.: MSUC 24 (2). Hickman Co.: MSUC 212 (1). BAYOU DE CHIEN SYSTEM. Kentucky: Hickman Co.: MSUC 420 (3), MSUC 423 (5), MSUC 424 (13), MSUC 495 (2), MSUC 426 (4). Tennessee River drainage —TENNESSEE RIVER SYSTEM. Alabama: Limestone Co.: UMMZ 115038 (1), UMMZ 115049 (1), UMMZ 115121 (1), UMMZ 122637 (6), UMMZ 122707 (2). Madison Co.: UMMZ 115189 (1). Morgan Co.: UMMZ 115403 (1). Misssissippi: Tishomingo Co.: UAIC 43859 (1), UAIC 4369 (1). Tennessee: Benton Co; UMMZ 168331 (1), UT 91.795 (4). Decatur Co; TU 91-812 (11) Sebiarcinn Gor: UMMZ 177666 (1), UT 91.249 (7). Henderson Co.: UT 9I26Rea wr 91.796 (11). Henry Co.: UT 91.883 (6). Humphreys Co.: UT 91.797 (13), TU 91.811 (18), UT 91.855 (18). Perry Co.; UW) 916231) Rake aiatelay: Marshall Co.: UMMZ Uncat. Coll. (1). Ohio River drainage -GREEN RIVER SYSTEM. Kentucky: Ohio Co.: USNM 117343 (2), USNM 63797 (2). (JDW) Office of Endangered Species, Fish and Wildlife Service, Depart- ment of the Interior, Washington, D.C. 20240; and (DAE) Department of Zoology, University of Tennessee, Knoxville, Tennessee 37916. PROC. BIOL. SOC. WASH. 90(1), pp. 19-29 APOGON MOSAVI, A NEW WESTERN ATLANTIC CARDINALFISH. AND A NOTE ON THE OCCURRENCE OF APOGON LEPTOCALUS IN THE BAHAMAS George Dale The fishes of the Bahamas are perhaps better known than those of any other tropical marine area of comparable size. Thirteen previously de- scribed species of the genus Apogon have been recorded in the Bahamas (Bohlke and Chaplin, 1968, with the generic reassignments prescribed in Fraser and Robins, 1970), making it the most speciose genus in the ichthyofauna of that area. Most, perhaps all, of the species occur through- out the Bahamas (and probably throughout much of the tropical western Atlantic) and thus may be considered broadly sympatric. These considerations have made the genus Apogon an ideal group in which to investigate the ecological principle of competitive exclusion. This paper is an outgrowth of that investigation. A new species of Apogon is described from the Bahamas, Haiti, and Jamaica, and Apogon leptocaulus Gilbert is recorded for the first time in the Bahamas. While teaching a marine biology course given in Nassau, Bahamas, in June, 1975, I spent some time observing the sea anemone association of Apogon quadrisquamatus reported by Colin and Heiser (1973), and made some collections. I became aware of certain differences between that species and some of the specimens identified as such that I had examined in the fish collections of the American Museum of Natural History, specifi- cally a large sample from Cat Island, Bahamas, collected in 1968 by C. Lavett Smith and James C. Tyler. Scrutiny of that sample revealed that it was indeed distinct from, though closely related to, A. quadrisquamatus. An additional specimen from the Bahamas, collected by C. L. Smith in 1966, two large samples, one from the Bahamas and one from Haiti, in the fish collections of the Academy of Natural Sciences of Philadelphia (ANSP), and a small Jamaican sample from the University of Miami Marine Laboratory (UMML) provided the study material. Meristic data in the description are based on 20 specimens, except lower limb gill raker counts (224 specimens) and preopercular margin serrae counts (42 specimens). Morphometric data are based on 12 specimens, except that the upper jaw length, ventral fin length, and caudal peduncle length, are based on measurements of 42 specimens. The holotype is deposited at the American Museum of Natural History (AMNH). Paratypes are deposited at AMNH, ANSP, UMML, and the National Museum of Natural History (USNM). Comparative material (A. quadrisquamatus ) was obtained from AMNH, ANSP, and USNM. Mea- 20 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON surements and counts were made according to Hubbs and Lagler (1958) and are presented in a format similar to that of Bohlke and Chaplin (1968). Apogon mosavi, new species Dwarf cardinalfish ieee Holotype —AMNH 35782 (29.7 mm SL), Bahamas, Cat Island, Hawks- nest Creek, 42 mi from shore, patch reef at dropoff, depth 85 to 105 ft, 31 January 1968, C. L. Smith and J. C. Tyler. Paratypes (collected with the holotype)—AMNH 34481 (111 specimens, 18.6-34.0 mm SL); ANSP 137482 (5, 26.7-28.1); UMML 32962 (5, 28.3- 29.5); USNM 216595 (5, 27.9-29.0). Other material examined—AMNH 29896 (1, 33.6 mm SL), Bahamas, Great Abaco, Southwest Point, 300 ft from shore, patch reef, depth 40 ft, 9 December 1966, C. L. Smith and B. Hanna; ANSP 124036 (117, 17.7- 27.9), Bahamas, Grand Bahama, ca. 2 mi SW of Deep Water Bay, depth 90-95 ft, 6 December 1969, W. A. Starck, J. D. Starck, and P. Hopper; ANSP 113434 (558, 14.6-28.4), Haiti, Gulf of Gonave, St. Marc Channel, off Mt. Rouis, depth 120-135 ft, 16 September 1967, J. C. Tyler, H. A. Feddern, T. Devany, and J. Durocher; UMML 31181 (13, 26.4-30.9), Jamaica, Dis- covery Bay, Buoy Reef, depth 120 ft, 29 June 1972, P. Colin. Comparative material (Apogon quadrisquamatus ).—Bahamas: AMNH 27123, 28737, 28756, 28825, 29891, 30513, 30523, 33109, 33144, 33807, 34340; AMNH uncat.: field no. GD73-11, GD75-1, GD75-3, GD75-4 ,GD75-5, GD75-7; ANSP 97014, 97017, 97019, 97022, 97023, 97026, 97027, 97029. Florida: USNM 107325 (lectotype), 108888 (paratypes), 213474, 213475. Puerto Rico: USNM 207020, 207021, 213603, 213604, 213605. Grand Cay- man: ANSP 102440. Lesser Antilles: ANSP 106536, 106537, 106545. Nic- aragua: USNM 179231, 213601. Colombia: ANSP 112685; USNM 213606, 213607, 213608, Acc. 254315. Surinam: USNM 186430. Brazil: USNM 213602. Diagnosis.—A species of the genus Apogon, subgenus Apogon (see “Re- marks” below), with 8 segmented anal rays (last ray composite); 2-5 median predorsal scales; 13-16 gill rakers on lower limb of first gill arch (rarely 13 or 16, modally 15); 16 circumpeduncular scales; rear margin of | fleshy posteroventral portion of free preopercular margin anterior to a | ventral continuation of the line formed by the vertical edge of the pre- opercular margin; coloration (in alcohol) pale; scattered, slightly enlarged | melanophores on cheek and behind eye, giving these areas a freckled ap- | pearance; a basicaudal bar; no markings on fins. Description—Dorsal rays VI-I, 9 (last ray composite); anal rays II, 8 | (last ray composite); pectoral rays 12; lateral-line scales to caudal-fin | VOLUME 90, NUMBER 1 \ Mii, . \ 1.7 mm SL, off Cat Island, Bahamas. ype, AMNH _ holoty . Zon MOSAvI Apo 21 22 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON base 24: scales above lateral line 2; scales below lateral line 6 or 7; median predorsal scales 2-5; scales around caudal peduncle 16; gill rakers on first gill arch 4-6 (mostly 5) + 13-16 (mostly 15, rarely 13 or 16); serrae along vertical limb of free preopercular margin 8-20, on 42 specimens 19.4-34.0 moma SEs Greatest depth of body contained 2.7-3 times in SL; length of caudal peduncle 1.1-1.3 times in greatest body depth, 3.4-3.7 times in SL; least depth of caudal peduncle 1.7-2.1 times in its length; head length 2.2-2.3 times in SL; eye diameter 2.7—3.1 times in head length; snout short, its length about 5 times in head length; end of snout bluntly rounded; dorsal profile of head convex from snout to origin of dorsal fin. First dorsal spine 1.8—2.1 times in length of second spine; second dorsal spine slightly longer than third; second dorsal spine thickest spine in fin; spine of second dorsal fin 1.1-1.3 times in second spine of first dorsal fin; ventral fins short, extending to anus or slightly behind; innermost ventral ray connected to body by membrane at a point %-% of the way out on its mesial branch; pectoral fins narrow and rather long, extending to above point along anterior third of anal-fin base; first anal spine short, its length 4-5 times in length of second spine; second anal spine about equal in length to spine of second dorsal fin; caudal fin forked, the lobes rounded. Upper angle of rear margin of maxillary at or slightly behind a vertical at rear margin of pupil; rear margin of maxillary slightly concave; upper edge of maxillary slipping up beneath suborbital when mouth is closed; no orbital or anterior orbital serrations; posterior margin of preopercle serrate, the number of serrations tending to increase with the size of the specimen; a rounded flap of skin at angle of preopercle, not extending beyond an imaginary ventral continuation of the line formed by the free posterior margin of the preopercle; free tip of opercular spine short, narrow, and sharp; posterior nostril a vertically elongate opening nearer to eye than to anterior nostril. Large ctenoid scales present on cheek and opercle; scales on body all finely ctenoid, including those on thorax and nape, except for a few an- teriorly on thorax and those before pectoral fin base; scales on nape ex- tending forward to occiput; fins naked except for small scales basally on caudal fin; lateral line complete; lateral-line scales similar in size to adjacent body scales. Tiny villiform teeth in bands on jaws, a few near front of upper jaw and laterally on lower jaw very slightly enlarged; vomer and palatine with irregularly uniserial villiform teeth; palatine sometimes lacking teeth; gill rakers on first arch long, longest gill filament in outer series contained 3-4 times in longest raker; tip of lower jaw extends more anteriorly than upper jaw. Caudal skeleton with three separate elements in hypural fan (hypurals VOLUME 90, NUMBER 1 23 1 and 2 fused, 3 and 4 fused, 5 separate) in 211 of 225 specimens X-rayed, four elements (hypurals 1 and 2 fused, 3, 4, and 5 separate) in 14 speci- mens; 10 precaudal + 14 caudal vertebrae; 2 predorsal bones located antero- dorsal to neural spines of first and second vertebrae, respectively. Color in life unknown; color in alcohol pale with slightly enlarged melano- phores scattered on cheek and behind eye, giving these areas a freckled ap- pearance; a dense pattern of small melanophores on nape make this region dusky; upper half of body has peppering of tiny melanophores, which tend to be concentrated on the edges of the scales; lower half of body with little or no pigmentation; dark brown bar at base of caudal fin about as wide as pupil; this bar usually slightly wider dorsally; the posterior margin of this bar just anterior to the posterior margin of the hypural fan; all fins color- less; a silvery iris, a silvery iridescence shining through integument of cheek and opercular region, and a silvery peritoneum shining through body wall are conspicuous on the holotype and the other specimens in the type col- lection, but rare or absent (probably an artifact of preservation) in the other collections; peritoneum often marked with scattered, enlarged melano- phores. Morphometric data for the holotype and 11 other specimens (4 para- types from AMNH 34481, 2 specimens each from ANSP 124036, ANSP 113434, and UMML 31181, and the single specimen AMNH 29896) are given in the following paragraph. The measurements given for each char- acter designate, in order, the holotype, the mean of all 12 specimens, and the range of all 12 specimens. All measurements are given in thousandths of SL. Greatest body depth 360, 348.8, 329-365; greatest body width 182. 178.8, 146-196; snout to first dorsal fin 458, 443.3, 424-460; snout to second dorsal fin 620, 622.7, 610-638; snout to anal fin 646, 645.3, 606-671; snout to pectoral fin 414, 424.1, 405-453; snout to ventral fin 407, 413.8, 393-464: snout to anus 630, 615.5, 587-636; least depth of caudal peduncle 162, 150.5, 135-162; caudal peduncle length 273, 278.2, 266-291; head length 451, 438.3, 428-451; head depth at occiput; 303, 301.3, 264-325; eye diam- eter 158, 149.8, 142-162: snout length 88, 88.3, 83-95; upper jaw length 292, 222.8, 214-935; bony interorbital width 91, 94.2, 85-103; first spine of first dorsal fin 91, 86.0, 71-92; second spine of first dorsal fin 182, 169.5, 155-186; spine of second dorsal fin 155, 149.2, 136-163; longest dorsal soft ray 256, 246.3, 234-262: first anal spine 37, 32.7, 27-37; second anal spine 148, 143.1, 132-160; longest anal soft ray 210, 205.4, 187-227; longest pec- toral ray 249, 244.3, 232-258; ventral spine 155, 150.6, 135-179; ventral fin length 215, 207.3, 198-219; caudal fin length 306, 315.8, 306-335; shortest caudal ray 239, 232.4, 219-240. Remarks.—Assignment of this species to Apogon is based on its close similarity to Apogon quadrisquamatus Longley. However, in contrast to other Apogon species, having five free hypurals, the new species (and ‘soATzeBOU ARI-X WOIF poledoid saatyisog “(TS WW 6'PE “Z-SLGD ‘ou perf “youn FANNY) snwjnopw ‘vy “dD “(TS wu TNE ‘T-GLGD ‘ou plelf “yeoun FINWY) snzwonbsiuponb “vy “gq “(TS ww LOS “T8PPE HNWY) tavsow uosody ‘y ‘speanddsy da1f QAIF SMOYS FZ ‘P pur |g °Z pure | sjeinddy Jo suoisny Moys g puke YW ‘sotoeds uosody 9ea14} JO suoJ[OYs [epneo [eoidAy, °% ‘SI PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 24 VOLUME 90, NUMBER 1 25 Table 1. Fusions of hypural bones in caudal skeleton of 225 specimens of Apogon mosavi (from 4 collections) and 104 specimens of A. quadrisquamatus (from 19 col- lections ), as determined from X-rays. A. mosavi A. quadrisquamatus 3 hypurals (1 and 2 fused, 3 and 4 fused, 5) 211 1a 4 hypurals (1 and 2 fused, 3, 4, and 5 separate) WAS 37 4 hypurals (3 and 4 fused, 1, 2, and 5 separate ) = 167 5 hypurals (no fusions) — 14+ * The separation of hypurals 3 and 4 in these 14 A. mosavi specimens is a minute but distinct separation of the posterior *%4 of the length of these bones. Anteriorly, a faint line indicates a continuation of the separation, but this is questionable. + “Non-fusions” in A. quadrisquamatus are typically rather wide separations, similar to those of other Apogon species. also A. quadrisquamatus) usually has three hypural elements (Table 1, Fig. 2). Nevertheless, all other characters studied place it in Apogon, and Fraser's (1972) keys to apogonid subfamilies, genera, and subgenera place it (and A. quadrisquamatus) in Apogon (Apogon). The diagnosis given above for A. mosavi distinguishes it from all other species assigned by Fraser to this subgenus, and in the western Atlantic from all other shallow water cardinalfishes (including Apogon (Apogon) species, Apogon of other subgenera, Phaeoptyx, and Astrapogon). The characters in the diagnosis which distinguish Apogon mosavi from A. quadrisquamatus, with which it would most likely be confused, are those involving pigmentation and gill raker counts. These, as well as cer- tain morphometric characters separating the species, are detailed below. In alcohol there are no dark markings on the head (ignoring the rather inconspicuous dusky region on the nape) except scattered melanophores, larger than those on the body, distributed in the region immediately below and behind the eye. In A. quadrisquamatus the head is peppered all over with small melanophores, similar in size to those on the body, and there is usually discernable a concentration of these into a slender triangular streak extending posteroventrally from the posteroventral border of the eye. The body of A. mosavi is pale, with pigmentation in the form of scattered, tiny melanophores usually limited to the dorsal halt of the body, Body pigmentation of A. quadrisquamatus is variable, sometimes dark, and occurs both dorsally and ventrally. A. mosavi lacks pigmentation on the fins, except for a few rare instances of a hint of pigmentation on the first dorsal fin, in contrast to A. quadrisquamatus, which nearly always has some pigmentation on the first dorsal fin. In some dark specimens this takes the form of an irregular streak across the second and third spines. Such specimens also tend to have a dusky stripe at both the second dorsal- fin base and the anal-fin base. A. mosavi has a basicaudal bar, nearly 26 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Table 2. Gill raker counts on lower limb of first gill arch, done on right side of 224 specimens of Apogon mosavi and 63 specimens of A. quadrisquamatus. Counts include raker at angle and all rudiments. Number of Lower Limb Gill Rakers 12 13 14 15 16 A. mosavi — 2 101 by 4 A. quadrisquamatus 8 48 7 — — always quite distinct, while A. quadrisquamatus typically has a smudgy spot in that region. In pale specimens of the latter species this spot is | faint or lacking. In a few very dark specimens the spot is prominent and vertically elongate, approaching a bar-like appearance, but these speci- | mens typically have distinct fin pigmentation, a distinct triangular streak | at the eye, and dense pigmentation dorsally and ventrally on the body, characters always absent in A. mosavi. Differences in gill raker count between A. mosavi and A. quadrisquama- tus are shown in Table 2. Morphometric characters helpful in distinguishing A. mosavi from A. quadrisquamatus are slight differences in upper jaw length, ventral fin length, and caudal peduncle length (Table 3). Since the latter two char- acters, in which the interspecific difference is greatest, vary inversely in the two species, combining them as “caudal peduncle length/ventral fin length” provides a character ratio which maximizes the separation of the two species (Table 3, Fig. 3). Ecology.—The type collection was part of a large, diverse rotenone sample made at 85-105 ft, from a patch reef at the dropoff edge, about %% mi off Cat Island, Bahamas. This sample included five other apogonid species (Apogon affinis, A. lachneri, A. phenax, A. townsendi, and Phaeop- Table 3. Comparison of certain morphometric characters in Apogon mosavi and A. quadrisquamatus. Values for mean, range, and standard deviation (SD) of upper jaw, ventral fin, and caudal peduncle lengths are in thousandths of standard length. Caudal peduncle length/ventral fin length (C.P.L./V.F.L.) is a simple ratio. A. mosavi A. quadrisquamatus (19.4-34.0 mm SL) (19.4-57.9 mm SL) N Mean _ Range SD N Mean Range SD Upper jaw length 42 999.4 211-949 6.4 46 236.0 221-248 5.6 Ventral fin length 42 208.3 170-226 10.1 56 240.2 216-269 10.2 Caudal peduncle length 42 277.5 259-293 8.8 of 257.4 . QV(=9635 ies (Gal eee AVA ese ADT lesa 7169 O08 56 1.07 0.92-1.26 0,07 VOLUME 90, NUMBER 1 27 55 50 (MM) pS wn & So 35 30 STANDARD LENGTH 25 20 1.0 1.1 1.2 1.3 1.4 1.5 1.6 CAUDAL Srp anae | VENTRAL LENGTH FIN LENGTH Fig. 3. Relationship of standard length to the ratio of caudal peduncle length to ventral fin length, in Apogon mosavi ( @) and A. quadrisquamatus (x ). tyx conklini) and 36 species of other fish families. The other four known collections of A. mosavi were also part of samples from coral reef habitats taken at depths of 40-135 ft. The sample including the single specimen of A. mosavi (AMNH 29896) also included four specimens of the closely related species A. quadrisquamatus. The fact that the largest individual in the five collections (S16 speci- mens) of A. mosavi is 34.0 mm SL (one of the paratypes) indicates that this is a diminutive species (the upper size range of all other western Atlantic Apogon species, some known from far fewer specimens, sub- stantially exceeds 34.0 mm SL). This is supported by the finding of 11 specimens (23.4-28.5 mm SL) from two collections (ANSP 113434 and ANSP 124036) which are ripe females, and 13 specimens (21.8-27.7 mm SL) from the same two collections with eggs or embryos in the mouth. Most or all of these specimens evidently represent oral brooding in- dividuals. Details of these findings will be published elsewhere. 28 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Etymology.—tThe trivial name mosavi, an acronym for Mount Saint Vin- cent, recognizes the College of Mt. St. Vincent’s annual summer course in marine biology, conducted in the Bahamas. The suggested common name, dwarf cardinalfish, refers to the small size of this species, as detailed above. Apogon leptocaulus Gilbert The description of Apogon leptocaulus Gilbert (1972) was based on single specimens from three localities, one off Florida and two in the west- ern Caribbean (British Honduras and Providencia Island, Colombia). Identification of a single specimen (AMNH 33179, 27.7 mm SL), collected by C. L. Smith and J. C. Tyler on 22 January 1968 off Southeast Point at Mayaguana Island, as A. leptocaulus extends the known range of this species to the Bahamas. It was part of a rotenone collection from a patch reef at 70-80 ft. This specimen essentially agrees in descriptive details, meristics, and morphometrics with the type specimens. In addition, I have been informed (P. Colin, pers. comm.) that a pub- lished photograph of A. leptocaulus (Colin, 1974) was taken in the Baha- mas, on the Exuma sound side of Eleuthera at about 80 ft depth. Acknowledgments Ernest A. Lachner, USNM, William F. Smith-Vaniz and James E. Bohlke, ANSP, and C. Richard Robins, UMML, loaned comparative material. Thomas H. Fraser of the Environmental Quality Laboratory, Port Char- lotte, Florida, and Nancy Neff, Michael Oliver, and Richard Vari of the AMNH reviewed the manuscript and gave many valuable suggestions. Sister Kathleen Tracey of the College of Mt. St. Vincent, Riverdale, New York, provided the opportunity to do field work in the Bahamas. Mitch Alvo and John Nicholas aided in field work. Les Kaufman made me aware of the UMML Jamaica sample and M. Norma Feinberg brought to my at- tention the specimen that proved to be A. leptocaulus. C. L. Smith, AMNH, gave permission to use material collected by him, as well as much helpful advice. This research was aided in part by funds from the City University of New York. For all of this assistance I am most grateful. Literature Cited Bohlke, J. E., and C. C. G. Chaplin. 1968. Fishes of the Bahamas and adjacent tropical waters. Livingston Publ. Co., Wynnewood, Pennsylvania. , and J. E. Randall. 1968. A key to the shallow-water west Atlantic cardinal- fishes (Apogonidae), with descriptions of five new species. Proc. Acad. Nat. Sci. Philadelphia 120( 4) : 175-206. Colin, P. L. 1974. Mini-prowlers of the night reef. Sea Frontiers 20(3):139-145. , and J. B. Heiser. 1973. Associations of two species of cardinalfishes VOLUME 90, NUMBER 1 29 (Apogonidae: Pisces) with sea anemones in the West Indies. Bull. Mar. Sci. 23(3) :521—-524. Fraser, T. H. 1972. Comparative osteology of the shallow water cardinal fishes (Perciformes: Apogonidae) with reference to the systematics and evolution of the family. Ichthyol. Bull. Rhodes Univ. 34:1-105. , and C. R. Robins. 1970. The R/V Pillsbury deep-sea biological expedition to the Gulf of Guinea, 1964-65. 18. A new Atlantic genus of cardinalfishes with comments on some species from the Gulf of Guinea. Stud. in Trop. Oceanogr. 4(2):302-315. Gilbert, C. R. 1972. Apogon leptocaulus, a new cardinalfish from Florida and the western Caribbean sea. Proc. Biol. Soc. Washington 85(36) :419-426. Hubbs, C. F., and K. F. Lagler. 1958. Fishes of the Great Lakes region. Rev. ed., Cranbrook Inst. Sci. Bull. 26:1-213. (Reprinted 1970, Univ. of Michigan Press, Ann Arbor. ) Department of Ichthyology, American Museum of Natural History, New York, New York 10024. Present address.—Department of Biological Sciences, Fordham Univer- sity, Bronx, New York 10458. PROC. BIOL. SOC. WASH: 90(1), pp. 30-42 DESCRIPTION OF COLOMASTIX JANICEAE N. SP., A COMMENSAL AMPHIPOD (GAMMARIDEA: COLOMASTIGIDAE ) FROM THE FLORIDA KEYS, USA Richard W. Heard and Daniel G. Perlmutter During November of 1968 and 1973 and June of 1970 more than 100 specimens of an undescribed commensal amphipod belonging to the genus Colomastix Grube, 1861, were collected from loggerhead sponges, Sphecio- spongia vesparia (Lamarck), in the lower Florida Keys. Additional speci- mens of this new species, collected from Dry Tortugas, Florida, were borrowed from the Division of Crustacea of the U.S. National Museum of Natural History. Until the recent description of the genus Yulumara Barnard, 1972, from Australia, Colomastix was the only recognized genus in the family Coloma- stigidae. Although there is little ecological information available on the family, its members are generally considered to be closely associated or symbiotic with sessile marine invertebrates, especially sponges. The phylo- genetic position of the Colomastigidae is still unsettled. Bousfield (1973) stated that the family was “apparently derived from stenothoid-like an- cestors, whereas Barnard (1974) tentatively linked it with the dexaminid families (Ampeliscidae and Dexaminidae). In his comprehensive monograph on the gammaridean Amphipoda, Stebbing (1906) recognized Colomastix pusilla Grube, 1961; C. brazieri Haswell, 1880; and C. hamifera Kossman, 1880, as the only valid known species of Colomastix. He stated, however, that C. hamifera might be a juvenile male of C. pusilla. In this same work Stebbing listed Cratippus tenuipes Bate, 1862; Cratippus crassimanus Heller, 1866; and Exungia stilipes Norman, 1869, as synonyms of C. pusilla. Walker (1909) recognized Cratippus as a synonym of Colomastix, but considered C. crassimanus (Heller) as valid, a designation not accepted by later authors. Since the work of Stebbing (1806), 10 species have been described: C. fissilingua Schellenberg, 1926; C. castellata K. H. Barnard, 1932; C. simplicauda Nicholls, 1938; C. magnirama Hurley, 1954; C. subcastellata Hurley, 1954; C. japonica Bulycheva, 1955; C. lunalilo J. L. Barnard, 1970; C. kapiolani J. L. Barnard, 1970; C. halichondriae Bousfield, 1973 and C. keiskama Griffiths, 1974. See Della Valle (1893), Stebbing (1906), Schellenberg (1926), Hurley (1954), J. L. Bamard (1955) for additional references to the early literature on Colomastix. More recent studies by Ledoyer (1968), Bellan-Santini (1972) and Bellan-Santini and Ledoyer (1973, 1974) give additional information on the distribution and ecology of Colomastix pusilla and C. fissilingua. Table 1 lists the species of Colomastix and the localities VOLUME 90, NUMBER 1 31 from which they were originally described and presents a comparison of several important taxonomic characters. Colomastix hamifera, a probable synonym of C. pusilla, is not listed due to lack of information on important characters (i.e., nature of inner plate of maxilliped). Specimens collected during this study were fixed in 10% formaldehyde solution and transferred to 70% ethanol within 48 h. Dissected parts were studied in temporary water or glycerine preparations or mounted on slides in Turtox water soluble mounting medium. Illustrations were made with the aid of camera lucidas. Colomastix janiceae, new species Figs. 1-4 Colomastix pusilla—Pearse, 1932 (in part). Diagnosis.—Relatively large species, female and male reaching lengths (excluding antennae) of 7.5 and 9.3 mm, respectively. Head of male much larger than that of female, length approximately equal to that of first 2 thoracic segments. Prominent supra-epistomal process present in both sexes, widely separated from epistome in mature male. Inner plate of maxilliped completely fused, acutely triangular. Dactyl of male gnathopod 2 reaching posteriorly less than % length of propodus (palm). Gills well-developed: in female those of gnathopod 2 reaching over 7% length of basis (second seg- ment) and those of legs 1-4 distinctly longer than basis. Long simple setae on dorsal surface of pleopod peduncular segments of adult male. Inner ramus of uropod | distinctly longer in male; subequal in female. Outer ramus of uropod 3 approximately * that of inner ramus. Telson sub- triangular, distal % of margin with 15-20 castellations. Description—Body subcylindrical, narrower in male than in female, flattened slightly dorsoventrally. Length of adult males 6.5-9.4 mm; adult females (i.e., marsupium formed) 5.5-7.5 mm. Head: Wider than long in both sexes. Adult male: enlarged, approxi- mately equal in length to first 2 thoracic segments. Female and subadult male: head and first thoracic segment subequal in length. Eyes with 16-20 facets; orange-yellow in life, pale yellow in specimens preserved in 70% ethanol. Rostrum and interantennal (subrostral ) process subequal and acute (Fig. 4K); ventral margins of interantennal process slightly raised pro- ducing a shallow medial depression. Sharp-edged interantennal ridge con- cave (Fig. 4A, B, K). Indentation of supraantennal ridge reaching no farther than middle of eye. Epistome well developed in both sexes. Promi- nent antero- (supra) epistomal process (or keel) present in both sexes, directly adjacent to epistome in adult female (Fig. 4B) and widely sepa- rated from epistome in adult male (Fig. 44) (due to elongation of head in mature male). 32 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 1. Colomastix janiceae. A, Adult male (9.3 mm), lateral view; B, Adult female (7.5 mm), lateral view; C, Adult male, dorsal view; D, Adult female, dorsal view. Scale = 1 mm. Antennae: Subequal in length; approximately % length of body in both sexes. Antenna 1: First peduncle segment approximately equal to second, third nearly *s length of second; ventral and inner ventral margin of third peduncular segment of male with relatively long, simple, spine-setae (Fig. 2B, E); ventral and inner ventral margin of third peduncular segment of female with short spines, each with a subapical sensory hair; flagella with 3 visible segments bearing long spine setae and sensory aesthetascs (Fig. 2E, F). Antenna 2: Fourth peduncle segment about %4 longer than third, length of third and fifth peduncle segments nearly equal; flagella with 3 visible segments, first much larger and longer than other 2 combined; spination of fourth peduncle segment and size of flagellum segment 1 differ with sex (Fig. 2G, H). See Fig. 2A-H for additional details on the spination, setation and morphology of antennae | and 2. VOLUME 90, NUMBER 1 33 Fig. 2. Colomastix janiceae. A, Ant. 1 2 inner view; B, Ant. 1 ¢ inner view, last ped. seg. showing long setae along ventral and inner ventral margin; C, Ant. 2 9@ inner view; D, Ant. 2 9 showing details of basal ped. segs.; E, Ant. 1 ¢ showing detail of last ped. seg. and flagellum (inner lateral aspect); F, Ant. 1 @ showing detail of last ped. seg. and flagellum (inner lateral aspect); G, Ant. 2 ¢ showing detail of ped. seg. and flagellum (ventrolateral view); H, Ant. 2 @ showing detail of last ped. seg. and flagellum (lateral inner view). Scale: A = 0.5 mm; B = 0.1 mm. Mouthparts: Mandibles small, ending in 5 strong spine-teeth, distal- most spine-tooth with trifid tip (Fig. 4F); molar process small, blunt, with concave medial face. Upper lip suboval, with fine short setae along rounded lateral and apical margins. Maxilla 1 (Fig. 4G) cheliform in appearance, relationship and origin of segments difficult to interpret, 3 or possibly 4 articles present; dactiliform (1 article) palp (?) opposing acutely tipped distal lobe of outer plate; inner plate (?) a small, finely setose lobe ( vesti- gial) located medially on inner face of basal segment (Fig. 4E). Mavxilla 2 (Fig. 4H) bilobed, smaller than maxilla 1, outer lobe narrower and longer than inner; both lobes with apical and subapical setae. Mavxilliped (Fig. 4T) forming opercular cap over other mouthparts; inner plate acutely triangular, entire (not distally cleft); outer plate with relatively straight distal margin and with 2 setae on lateral margin (inner noticeably larger than outer); palp with 4 segments; inner distal margin of segment 2 with a single seta; inner margin of segment 3 pubescent, with 2 strong proximal setae and single small more distal seta, single distal setae on outer margin 34 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 3. Colomastix janiceae. A, Distal end gnath. 2 of 6.5 mm ¢, outer aspect; B, Gnath. 2 of 6.5 mm 64, inner view of last 3 segs.; C, Gnath. 2 large (9.3 mm) ¢, inner view; D, Gnath. 1 of 6.5 mm ¢; E, Gnath. 1 large (9.3 mm) ¢; F, Gnath. 1 adult 2; G, Gnath. 2, coxal plate and gill, adult 9; H, Pereopod 2, coxal plate, oostegite, and gill of adult @; I, Pereopod 4, coxal plate and gill of adult 2; J, Telson of adult 9°, dorsal view; K, Ventral view of abdomen of 6.5 mm ¢ showing penal organs at base of 5th pereopod; L, Pleopod 3 of adult 2 (ant. aspect); M, Pleopod 3 of 9.3 mm 6 (ant. aspect); N, Ul of adult 9; O, U2 of adult 9; P, U3 of adult 9. Scales: a= 0.5 mma | Bags. (A! eb ==3120 mm (hig. K) c= 0.o.anmA(. Figs. [ak =O): OE EE Ee VOLUME 90, NUMBER 1 35 Fig. 4. Colomastix janiceae. A, ventrolateral view of head and mouthparts, 9.3 mm 6; B, Same, adult 9; C, Ventral view of head and mouthparts, adult 2; D, Same, 9.3mm ¢; E, Ventral aspect max 1, max 2, upper lip, and epistome of adult @; F, Two ventral views of mandible of adult 9; G, Max 1 of adult 9; H, Max 2 of adult 9: I, Mxpd of adult 2; J, 6.5 mm 4, lat. view of anterior end; kK, Adult 9, lateral view of anterior end. Scale: a = 0.5 mm (Figs. A, B); b = 1.0 mm (Figs. C, D, J, K); e = 0.2 mm (Figs. E-H); d = 0.5 mm (Fig. I). at joint with dactyl; segment 4 (dactyl) over “ length of segment 3, inner margin finely pectinate. Mouthparts of adult male smaller than those of adult female (Fig. 4A-D). Gnathopod 1—adult male: Atrophied, greatly reduced in size ( Fig. 4D); 36 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON propodus with small dactyl and 1-6 distal spines (Fig. 3D) [large adult male (9.5 mm) with only 1 small blunt distal spine (Fig. 3E), smaller adult male (6.5 mm) with 6 distal spines, 2 more strongly developed than others (Fig. 3D)]; propodus longer than carpus. Subadult male: re- sembling that of female. Female (adult and subadult): not reduced in size; propodus with distal brush of 7-8 curved compound setae and elon- gate, spine-like dactyl (Fig. 3F ); propodus slightly shorter than carpus. Gnathopod 2—adult male (Fig. 3A-C): Well developed; propodus (palm) with 3 strong distoventral teeth opposing dactyl; dactyl reaching posteriorly less than % length of palm, a blunt low tooth or tooth-ridge on mid-ventral margin in large adult male; ventral margin of carpus rounded, not acutely produced; distal portion of basis dilated with rounded margins (Fig. 3C). Subadult male: similar to that of adult female. Adult and sub- adult female with simple or marginally subchelate (Fig. 3G) propodus and carpus subequal in length. Setation of gnathopod 2 (both sexes) as illustrated (Fig. 3A—C, G); part of ventral inner faces of propodus, carpus and merus pubescent in adults of both sexes (Fig. 3C, G). Pereopods: Typical of genus, 1-2 longer than 3-5; 3-5 with orientation reversed; distal end of article 2 (basis) of pereopod 5 wider than that of pereopod | or 2; article 6 (propodus) longer than article 4 or 5, article 4 longer than article 5. Coxal plates (Figs. 1; 4J, K): Strongly developed; 2-7 with medial longi- tudinal ridge, more strongly developed on plates 2-5, more prominent on adult female than on male and subadult; anterior margins of plates 2-5 over- lapping posterior margins of preceding plates, anterior margins of plates 6-7 covered by posterior margin of preceding plates; anterior margin of plates 2-4 in adult male and 2-5 in adult female projected forward forming a blunt process, more prominent in adult female. Pleopods: Peduncular segments stout, equal in width; those of pleopod 1 slightly longer than those of second and about % longer than those of third pair; width-length ratios (from anterior to posterior) 4:11, 2:5 and 1:2; inner distal margin of each segment armed with pair of coupling setae (Fig. 3L); segments 1-3, respectively, in adult male, with 0-2, 20-23 and 7-9 long simple setae near margin on dorsal surface near inner distal margin (Fig. 3M); no such setae on peduncle segments 1-3 of female. Rami equal in length with 4 articles, article 1 longer than distal 3 articles combined; inner medial margin article 1 (anterior to posterior) with 3-4, 3, and 2-3 compound setae, respectively (Fig. 3L, M); rami (outer and inner) with long compound swimming setae, 1 pair originating at distal end of each article; rami (excluding setae) distinctly shorter than peduncle segments. Uropods: Rami (both sexes) lanceolate with margins finely pectinate, tips of U2 reaching nearly to tips of U3. Female: rami of U1 narrow and lanceolate, equal in length and slightly shorter than peduncle; peduncle VOLUME 90, NUMBER 1 37 % longer than that of U2 and twice the length of peduncle U3. Outer ramus of U3 narrower and slightly shorter than inner, length of inner ramus nearly equal to peduncle length. Inner ramus of U3 robust, % longer than outer peduncle, width over % length; peduncle and outer ramus nearly equal in length. Male: outer ramus of Ul much narrower and nearly % shorter than that of inner. U2 and 3 similar to those of female. Telson (Fig. 3J): Subtriangular, distal % nearly triangular with 15-20 relatively inconspicuous castellations; 3-4 pairs of inconspicuous small setae on dorsal surface near margins, | minute pair at tip. Female: width approxi- mately % length. Male: narrower with tip slightly turned up. Holotype.—Adult female, USNM No. 152666. Paratype.—Adult male, USNM No. 152667. Type-locality Molasses Keys (Florida Keys), 27 November 1969, depth earn: Other localities —Key West and Dry Tortugas, Florida. Habitat—In sponges, principally Spheciospongia vesparia (Lamarck). The species is named in honor and memory of the late Edna Janice Heard, a beautiful and rare person. Comparisons and Discussion Colomastix janiceae, C. pusilla, C. japonica, C. lunalilo, and C. keiskama are the only described species reported to have the inner plate of the maxil- liped entire or completely fused. The condition of the inner plate in C. brazieri and C. halichondriae was not determined; however, we examined specimens of C. halichondriae from Georgia (USA) and found the inner plate to be completely fused. In another species, C. magnirama, only a shallow cleft is reported present on the distal tip of the inner plate. The structure of the telson, the greatly enlarged inner ramus of uropod 3, and the pronounced carpal tooth on pereopods 3-5 further distinguish C. magnirama from C. janiceae. The similarity of gnathopod 1 in both sexes, and the outer ramus of uropod 3 being half or less as long as the inner, separate C. brazieri, C. japonica, and C. lunalilo trom C. janiceae. The much smaller body size, gnathopods 1 being similar in both sexes, and the shape and relatively smooth margin of the telson distinguish C. halichondriae and C. keiskama from C. janiceae. Of the described species, Colomastix janiceae is most similar and apparently most closely related to C. pusilla (sensu stricto). The gnathopods of C. pusilla and C. janiceae have the same general morphology. In the adult male, gnathopod 1 of both species is greatly reduced in size (vestigial); however, C. pusilla has much smaller gills, a more finely castellate (or serrate) telson margin, and the rami of uropods | and 3 are nearly equal in both sexes. Like C. pusilla and C. janiceae, the “C. pusilla” of J. L. Barnard (1955) from Hawaii has gnathopod 1 greatly reduced in the adult male. Based vsn IOUUL (OLGT ) pavusreg "y “[ TIVMVPT & Ol] a SOYOIOU [VISIP G sndivo 0} ah fc) (oat si | wupjoidpy * puv[vo7, IOUUT S9YO}OU 10 SUOTILT (PS6T ) Aopnyey MON & Oxl] a) -[99S89 O8.1e] YIM sndieo 0} ahi) (oyun Colts DID]JAISDIGNS *D puryleo7 IQUUL dij 38 suoye} snpodo.d jo yoyo (PS6T ) Aopnyzy MON & OX] a) “UspUl MOTTVYS ZS YBuaT > =Apysys gE DUDMUSDUL * BOLIFY *§ “4seiy IQuUl snpodo.d jo 6L) syaggirg = gourtAoig odv & OXI % x yqoous = yyBuey] 54, S o.1}UO ; DUDYSIOY * ELON) O AH p I I [“% OV PRCA [2D Apnyjs juoso.id VSN jo OUUL snpodo.d jo ‘(EL6L ) Pleysnog JSVOD Jey & OAL] %s yoows = yysuoy % ~ aljuo = (y'E aDIupuoyoyvy *D uede[ IOUUL snpodo.d jo (SG6T ) BAdYo[Ng Jo BIS & OXI] i yjoows Yysus] % > alqua = G'G powodpl + vsn IQUUL snpodo.d jo (OLGT ) pleuteg Ty ‘f TIVMVET & oy] a oous = Ys us] 4 > ol1quo GT oyun) *O VSN JOUUL poyo}0U LO oyeTJoysvo — snpodo.id jo Apnjs yueso.id ‘SsAOY VPLlOp LT [BISHSOA ue a Ajouly Yj; [esIp —- YSU] > oud £"G apaaupl *y Apnijs yuosoid ‘(S68T ) AeA BI[Pa Aye Vy oyeppeysvo AjpourF.to — snpodo.d Fo ‘(GZG6T ) 98k YT Y xNoIADY) ‘pode Ny [VIsQseA Jou > a7VV.Llos Ve [eISIP Ysus] 4 > olqua—()'G pyisnd * 9 rr ennnnnnes PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 38 SODUDLIFOY, AU[CIOT ) eee En UOS|O} oD) 9vur poedy (urur) soloods od J, ype | SnUIe.L jo JO [AJoRp Jo -[Exeul Ys] podoyjyeus 10}no UIBIBIA UWOISUd}xO jo Apoq quout = =—- Ys U'T pleayorg oyefd =uMouy -doJaaoc{ Jouuy = wim ut “Txt ssn ‘XUSDWULOJOD, FO soloods ET Jo uostieduoy “T 9[qey 39 _—_———— eee UAZDLY “YD DPNDIVAUMS °F DnGUIpssif “Dd DID]JAISVI *D setoedg eITe.4sny IouUl petodet snpodoid yo poysoda. (OSS8T ) [Josey ‘uosyor{ wOg = =(¢)d Oy] qa jou §=6ysua 5 100 > *OT= SoLG “Ao8ST Spurysy Iouul (SS6T ) STTOYDIN ouienboryy = uAouyuN a yqoous uMouyun yoyo Gg S.0€ 009 “MoSP (SE6L) preureg "FRY spurs] TOUUI snpodo.d jo (9Z6T ) Stoquapeyos AOUYIO °*S & Oy] a soyojou [e}sIp GZ sus] YS wp OF So€S “Mo09 spurs] TOUUL snpodo.d jo (ZE6T ) preureg ‘y-y puepye db Xl] a oyel[9seO YSU] S > WP OF a ee ee a ee sooualoyoy AFTTRIOT eyeul en uos|e} GD) 2[eul Fo pedi (wut) ad J, ype T SnUIeLI jo [Aqep Fo -[rxeul = -Yysuey] podoyjeus 1ayno UIBAR IA UOISUd}Xx9 jo Apoq yuoul yisue'T plemyoeg oyejd = uMoUy -do]aaoq Jouuy = uImnut EIN VOLUME 90, NUMBER 1 ‘ponunuoy —— 555 ee eS eee Se HOS) ialL, 40 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON on Barnard’s description and illustrations, however, the Hawaiian specimens have a non-castellate or non-serrate telson margin and a relatively weakly toothed palm on gnathopod 2 of the male. If Barnard’s observations were correct, the Hawaiian form probably represents a distinct species. As men- tioned by Schellenberg (1926), some of the earlier reports of C. pusilla, especially those from areas other than the Mediterranean, may represent records of distinct and as yet unnamed species. For example, Bonnier’s (1893) specimens of “C. pusilla,” due to their uniquely shaped telsons (see Bonnier’s figure), appear to fit into this category. We examined several lots of specimens from Dry Tortugas (Florida) identified as C. pusilla by Pearse (1932) and Shoemaker (unpublished records). This material con- tained specimens of C. janiceae, and those of two additional new species to be described later. One of these new forms appeared similar to the material previously reported as “C. pusilla” from the northern Gulf of Mexico by Pearse (1912) and the other, a small species (2 mm) is characterized by the male having the inner ramus of uropod 1 greatly enlarged and curved dorsally. In addition to Pearse’s reports (1912, 1932) there are two other pub- lished records of “C. pusilla’ from the Northwestern Atlantic. Based on a single male specimen, Kunkel (1910) reported C. pusilla from Bermuda. Examination of the description and figure indicates that the Bermuda speci- men has closer affinities with C. pusilla than with C. janiceae. Additional material from Bermuda will have to be examined before any firm con- clusions can be drawn. Shoemaker (1942) identified a female specimen, collected from Isla de Providencia in the Caribbean, as C. pusilla. The fact that C. janiceae and at least 2 undescribed species of Colomastix from the Caribbean and adjacent areas have been confused with C. pusilla pre- viously (Pearse, 1912, 1932; Shoemaker, unpublished records) makes Shoe- maker's 1942 record questionable. Although C. hamifera and C. crassicornis, described from the Red Sea and the Adriatic, respectively, have been con- sidered synonyms or possible synonyms of C. pusilla (Della Valle, 1893; Stebbing, 1906; Hurley, 1954), such designations should be only tentative pending studies of additional specimens from their type-localities. Specimens of Colomastix janiceae occurred in all of the 15 Spheciospongia vespara we examined. Eight of these sponges were from Molasses Keys and 7 were from Key West; all were collected from depths of less than 2 m. In the Florida Keys we observed the following animals associated with C. janiceae in the canals of S. vespara: Synalpheus brooksi Coutiére, Syn- alpheus pectinger Coutiere, Typton tortugae McClendon, Balanus declivis Darwin, Nebalia cf. bipes (Fabricius), Leucothoe spp., Trypanosyllia zebra (Grube), and large numbers of harpacticoid copepods. In general these animal associates are similar to those reported by Pearse (1932) from S. vespara at Dry Tortugas. In 2 studies on the biocoenoses of S. vespara and VOLUME 90, NUMBER 1 4] other sponges from the Bahamas, Pearse (1950) and Jaronski (1969) did not report finding specimens of Colomastix. Acknowledgments We wish to thank J. L. Barnard and E. L. Bousfield for their constructive comments on the manuscript. Thomas E. Bowman and J. L. Barnard loaned specimens from the United States Museum of Natural History and Gertraud Krapp kindly supplied specimens of Colomastix pusilla from Italy. We would also like to acknowledge Walter B. Sikora and John Crawford for helping to collect specimens. During part of the study, lodging and labora- tory space were made available by the Pigeon Key Field Station of the Uni- versity of Miami. The senior author wishes to express his appreciation to A. G. Fish and R. M. Overstreet for their encouragement and assistance. Literature Cited Barnard, J. L. 1955. Gammaridean Amphipoda (Crustacea) in the collections of the Bishop Museum. Bernice P. Bishop Mus., Bull. 215:1—46. 1970. Sublittoral Gammeridea (Amphipoda) of the Hawaiian Islands. Smith- sonian Contrib. Zool. 34:1—-286. 1971. Keys to Hawaiian Marine Gammaridea, 0-30 meters. Smithsonian Contrib. Zool. 58:1—135. 1972. Gammaridean Amphipoda of Australia, Part I. Smithsonian Contrib. Zool. 103:1-333. —. 1974. Evolutionary patterns in Gammaridean Amphipoda. Crustaceana 27 (2): 137-146. Barnard, K. H. 1932. Amphipoda. Discovery Reports 5:1—326. Bellan-Santini, D. 1972. Amphipodes des milieux Portuaires. Tethys 3(2):255-263. , and M. Ledoyer. 1973. Inventaire des amphipodes gammariens récoltés dans la région de Marseille. Tethys 4(4):899-934. 1974. Gammariens (Crustacea—Amphipoda) des iles Kerguelen et Crozet. Tethys 5(4):635-708. Bonnier, J. 1893. Les amphipodes du Boulonnais. Bull. Sci. France Belgique 24: 161-207. Bousfield, E. L. 1973. Shallow-water gammaridean Amphipoda of New England. Cornell Univ. Press, Ithaca, 312 pp. Bulycheva, A. I. 1955. New species of Amphipoda gammarids from the Sea of Japan. II. Trav. Inst. Zool. Acad. Sci. URSS. 21:193-207. [In Russian. ] Chilton, C. 1912. The Amphipoda of the Scottish National Antarctic Expedition. Trans. Roy. Soc. Edinburgh 48(2) (No. 23):455-520, pl. 1-2. 1921. Report on the Amphipoda obtained by the F. I. S. “Endeavour” in Australian Seas. Biol. Res. “Endeavour” 5( 2) :32-92. Della Valle, A. 1893. Gammarini del Golfo di Napoli. Fauna und Flora des Golfes von Neapel und der Angrenzenden Meeres—Abschnitte, Monogr. 20, 948 pp. Griffiths, C. L. 1974. The Amphipoda of Southern Africa. Part 4. The Gammaridea and Caprellidea of the Cape Province east of Cape Agulhas. Ann. S. African Mus. 65(9):251-336. Grube, A. E. 1861. Ein Ausflug nach Triest und dem Quarnero. Beitrage zw Renniniss Thierwett dieses Gebietes ( Berlin). 42 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Haswell, W. A. 1880. On some additional new genera and species of amphipodous crustaceans. Proc. Linn. Soc. N. South Wales 4:319-350. Hurley, D. E. 1954. Studies on the New Zealand Amphipodan fauna. No. 6. Family Colomastigidae, with descriptions of two new species of Colomastix. Trans. Roy. Soc. New Zealand 82(2):419—429. Jaronski, S. T. 1969. Compositional variations in sponge biocoenoses. Yale Scient. Mag. 43(7):18-21. Kossman, R. 1880. Malacostraca. Zool. Ergebn. Reise in die Kiistengebiete des Rothen Meeres, II. Halfte, I. Lfg., III. Teil, Leipzig, 64-140, pls. 4-15. Kunkel, B. W. 1910. The Amphipoda of Bermuda. Trans. Connecticut Acad. Sci. IGG, fee, ey Ledoyer, M. 1968. Ecologie de la vagile des biotopes Mediterranéens accessibles en scaphandre autonome (Region de Marselle principalement). IV. Synthése de létude écologique. Rec. Trav. Sta. Mar. Endoume 56 (Bull. 40) :103-149. Nicholls, G. E. 1938. Amphipoda Gammaridea. Sci. Rep. Australian Antarctic Exped. 1911-1914. Ser. C, Zool. & Bot. 2(4):1—-145. Pearse, A. S. 1912. Notes on certain amphipods from the Gulf of Mexico, with de- scriptions of new genera and new species. Proc. U.S. Nat. Mus. 43:369-379. ——.. 1932. Inhabitants of certain sponges at Tortugas. In Papers from the Tortugas Laboratory. 7. Camegie Inst. Wash. Pub. 435:119-123. —. 1950. Notes on the inhabitants of certain sponges at Bimini. Ecology 31(1): 149-151. Schellenberg, A. 1926. Die Gammaridea der Deutschen Siidpolar—Expedition 1901- 1903. Deutsche Siidpolar-Exped. 28, Zool. 10:235-—414. Shoemaker, C. R. 1942. Amphipod crustaceans collected on the Presidential Cruise of 1938. Smithsonian Misc. Coll. 10(11):1-52. Stebbing, T. R. R. 1906. Amphipoda Gammaridea. Das Tierreich 21:1—806, figs. 1-127. Walker, A. O. 1909. Amphipoda Gammaridea from the Indian Ocean, British West Africa, and the Red Sea. Trans. Linn. Soc. London, Zool. (Series 1), 12(4): 323-344, plates 42, 43. (RWH) Gulf Coast Research Laboratory, Ocean Springs, Mississippi 39564; and (DGP) Skidaway Institute of Oceanography, Savannah, Georgia 31406. PROG. BIOL; SOC. WASH. 90(1), pp. 43-48 SARSIELLA PSEUDOSPINOSA, A NEW MARINE OSTRACOD (MYODOCOPINA; SARSIELLIDAE) FROM SOUTHERN CALIFORNIA James H. Baker From 1956 to 1960, the Allan Hancock Foundation of the University of Southern California undertook an oceanographic survey of the area from Point Conception to the Mexican border and extending seaward to the 100-m contour. This area is known as the mainland shelf portion of the con- tinental shelf off southern California. Of 1,498 stations occupied, 455 con- tained a total of 22 cypridinacean species (Baker, Ph.D. dissertation, 1975), including the following 3 species of Sarsiella: Sarsiella zostericola Cushman, 1906, known from the Atlantic and Pacific coasts of the United States, the Gulf of Mexico, and recently, England (Kornicker, 1975), was found in this study off Dana Pt., Newport Beach, Pt. Fermin, and Santa Monica. Sarsiella tubipora Darby, 1965, known from off the coast of Georgia, was found in southern California off San Diego, Oceanside, San Mateo Pt., Huntington Beach, Seal Beach, Man- hattan Beach, Venice, Santa Monica, Pt. Mugu, Pt. Las Pitas, Sand Pt., Loon Pt., Goleta Pt., Coal Oil Pt., Lento, and Pt. Conception. Sarsiella pseudospinosa, the new species, is described below. Order MYODOCOPIDA Sars, 1866 Suborder MYODOCOPINA Sars, 1866 Family SARSIELLIDAE Brady and Norman, 1896 Sarsiella pseudospinosa, new species Figs. 1-2 Holotype —AHF 5816, adult female from Station 5562, length 1.38 mm, height 1.15 mm. Valves in alcohol, appendages on slide. Type-locality —Station 5562, off Pt. Conception, California, 34°26’47’N, 120°21’35’W, bottom depth 19.2 m, water temperature 15.5°C, salinity 33.6%, Secchi disc reading 9.1 m. Material_—tThe holotype and a second female from Station 5631, off San Mateo Pt., 33°22/50’N, 117°34’55’ W, bottom depth 17.1 m, salinity 33.6 Secchi disc reading 3.0 m, sediment calcium carbonate 21.9%, sediment nitrogen 0.069%, in silty-sand. The second individual is in poor condition and, therefore, is not designated as a paratype. Etymology—Named for the spinose appearance of tube-pores covering the surface of the carapace. Remarks.—The holotype had a Foraminifera of the family Lituolidae attached to the carapace. a4 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON N A : A ATTN Ste ETT \ SFR « Sy ‘ 1 Be 11 ae My ox us mat \\ VLAN RE RR NN) ° 85 b Fig. 1. Sarsiella pseudospinosa, holotype, adult female: a, Entire view, lateral surface of right valve; b, Muscle scar pattern, medial view of left valve; c, Part of reticulated pattern, medial view of left valve; d, Infold area of rostrum; e, Infold area of caudal process. Description of Female.—Shell (Fig. la): Oval, greatest height anterior to middle; anterior produced in area of rostrum, incisur shallow but distinct, ventroposterior acuminate to form caudal process. Ornamentation (Fig. lc): Dorsal and ventral ridges, just within mar- gin, bearing 4 or 5 nodes; central nodose ridge, open anteriorly; surface with shallow reticulations; edge of reticulations, inter-reticulated area, margin, and ridges with hallow tube-pores, appearing to be spines. Infold (Fig. ld, c): Four short bare bristles, in a row, perpendicular to long axis of caudal process in right valve, only 3 in left valve; 2 hirsute bristles dorsal to caudal process; 2 very short bristles along inner margin of infold in front of caudal process. VOLUME 90, NUMBER 1 45 Fig. 2. Sarsiella pseudospinosa, holotype, adult female: a, Ist antenna; b, 2nd antenna, exopodite; c, 2nd antenna, protopodite and endopodite; d, Mandible; e, Maxilla; f, Maxilla, endites II and III enlarged; g, 5th limb; h, 6th limb: i, 7th limb; j, Furca. (B-basale, C-coxale, En-endopodite, Ep-epipodite, Ex-exopodite, P-protopo- dite ). Selvage: Narrow with wide lamellar prolongation with smooth outer margin. Muscle scars (Fig. lb): Circular pattern of 6, ovoid, some divided, ventroanterior to midpoint. Pore canals: Normal pores, hollow, tubular spinose; radial pores with long setae; setae distributed from anterior ventrally to caudal process. First antenna (Fig. 2a): Ist podomere bare; 2nd podomere with | spinose dorsal bristle; 3rd podomere with 1 ventral and 1 dorsal bristle: 46 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Table 1. Comparison of females of North American species of the Sarsiella carinata group (Kornicker and Wise, 1962). Carapace length No. bristles No. claws Species (mm ) 7th limb furca Sarsiella pseudospinosa Baker 1.38 6 terminal 5 2 lateral Sarsiella tubipora Darby 1e55 6 terminal 5 2 lateral Sarsiella spinosa Komicker and Wise 1.43 4 terminal 5 3 lateral 4th podomere not separated from 3rd, 1 dorsal bristle, 2 long ventral bristles; 5th podomere with 1 stout ventral sensory bristle; 6th podomere with 1 medial bristle; 7th podomere with a-, b-, and c-bristles; 8th podomere with d-, e-, f-, and g-bristles, bare, about same length as sensory bristle of 5th podomere. Second antenna (Fig. 2b, c): Exopodite of 9 podomeres; 2nd-8th with 1 stout bristle with hairs along both margins and minute spines along ventral margin; 9th podomere with similar bristle plus 1 short bare bristle. Endo- podite of 1 segment, bulbous, with 1 short distal chitinous protuberance, 2 short stout dorsal bristles. Mandible (Fig. 2d): Coxale short spine proximally and hairs along distal part of ventral margin; basale with 3 short stout ventral bristles. Endo- podite, Ist podomere with minute dorsal terminal spine and long stout terminal ventral claw; 2nd podomere same as Ist; 3rd podomere with 1 minute dorsal spine and long stout terminal claw. Maxilla (Fig. 2e, £): Exopodite with 1 long and 1 short bristle; basale with 1 short ventral bristle; lst to 3rd endites with 4 to 6 bristles. Endopo- dite, lst podomere with terminal spinous alpha and beta bristles; 2nd podomere with 2 a-bristles, 1 short c-bristle; 5 terminal stout b- and d- bristles with marginal denticles. Fifth limb (Fig. 2g): One endite with 1 short bristle. Exopodite, Ist podomere with 2 terminal bristles; 2nd-5th podomeres not separated, 2nd podomere with 3 bristles, 3rd-5th with 5 terminal bristles and 1 medial bristle; surface of 2nd-5th podomeres covered with fine hairs; epipodite with 20 hirsute bristles. Sixth limb (Fig. 2h): One endite with 2 spinose bristles; end podomere anteriorly with 2 rows of 13 bristles, 6 in one row and 7 in other, posteriorly with 2 long hirsute bristles, fine hairs along anterior and posterior margins. Seventh limb (Fig. 2i): Terminal end with 6 bristles, 3 on each side; 2 proximal bristles, 1 on each side; all bristles with 4 to 7 bells. VOLUME 90, NUMBER Il AT Furea (Fig. 2)): Each lamella with 5 claws, claw 4 broken off: claw 1 continuous with lamella, remaining claws separated by suture; claws 1-3 with scattered teeth and short hairs along posterior margin; scattered minute spines along posterior margins of lamella. Comparisons.—Long hollow tube pores resemble those of Sarsiella tubi- pora. However, in S. pseudospinosa, they differ by being about 2 to 3 times as long as in S. tubipora and closely resemble short spines. Sarsiella pseudospinosa also has the dorsal and ventral ridges in addition to the central nodose ring, which differentiates it from S. tubipora and S. spinosa Kornicker and Wise, 1962. It is a member of the S. carinata group of Kornicker and Wise (1962) and is compared to the North American mem- bers of the group in Table 1. Acknowledgments The samples containing the Ostracoda were kindly loaned to the author by Dr. John S. Grath, Allan Hancock Foundation. This research was sup- ported in part by a Grant-in-Aid of Research from the Society of The Sigma Xi and The University of Houston Coastal Research Center. Dr. Louis S. Kornicker critically reviewed the manuscript. The manuscript was typed by Mrs. Jeanne N. Rossman. Literature Cited Allan Hancock Foundation. 1965. An oceanographic and biological survey of the southern California mainland shelf. Appendix—Data. Califormia State Water Quality Control Bd. 27 (Appendix) :1—445. Baker, J. H. 1975. Distributions, ecology, and life histories of selected Cypridinacea (Myodocopida, Ostracoda) from the southern California mainland shelf. Ph.D. Dissertation, University of Houston, Houston, Texas. xivv + 185 pp. 27 figs. Brady, G. S., and A. M. Norman. 1896. A monograph of the marine and fresh water Ostracoda of the North Atlantic and of north-western Europe. Part IL, Sec- tions II-IV: Myodocopa, Cladocopa, and Platycopa. Sci. Trans. Roy. Dublin SOCmoeI 24.0. 021—7171 2-162) pls. Cushman, J. A. 1906. Marine Ostracoda of Vineyard Sound and adjacent waters. Proc. Boston Soc. Nat. Hist. 32:359-385. Darby, D. G. 1965. Ecology and taxonomy of Ostracoda in the vicinity of Sapelo Island, Georgia, pp. iii-vi, 1-76, text-figs. 1-89. Report No. 2 in: R. V. Kesling, D. G. Darby, R. N. Smith, and D. D. Hall, Four reports of ostracod investigations conducted under National Science Foundation Project GB-26. University of Michigan, Ann Arbor. Kornicker, L. S. 1965. Spread of ostracods to exotic environs on transplanted oysters, pp. 129-139. In F. M. Swain, L. S. Komicker, and R. F. Lundin (eds.), Biology and Paleobiology of Ostracoda. Bull. Amer. Paleontol. 65(282):1—G87. Paleon- tological Research Institution, Ithaca. —-— and C. D. Wise. 1962. Sarsiella (Ostracoda) in Texas bays and lagoons > Crustaceana 4(1):57-74. 48 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Sars, G. O. 1866. Oversigt af Norges marine Ostracoder. Norske Forhandl. Vidensk.- Selsk. Cristiania (1865)8:1-131. Department of Biology, University of Houston, Houston, Texas 77044. Based on material submitted in partial fulfillment of the requirements for the Ph.D. degree. Present address.—Southwest Research Institute, 3600 Yoakum Boulevard, Houston, Texas 77006, U.S.A. PROC. BIOL. SOC. WASH. 90(1), pp. 49-54 DESCRIPTION OF A NEW SUBSPECIES OF PRAIRIE VOLE, MICROTUS OCHROGASTER William D. Severinghaus The individual, age, sexual, and geographical variation in voles of the subgenus Pedomys has recently been analyzed as part of a monographic study of that group. Nearly 5,000 specimens from over 950 localities were examined for the following: dorsal and ventral pelage coloration; external measurements; cranial measurements (condylobasilar length, condylopalatal length, length of diastema, alveolar length of maxillary toothrow, palatilar length, length of incisive foramina, zygomatic breadth, lambdoidal breadth, prelambdoidal breadth, interorbital breadth, depth of braincase, and _pat- tern of M3); pelvic girdle parameters (ischial length, pubis length, pubis width, and shape of the descending portion of the ischium); hyoidal parameters (shape and size of the basihyal, hypohyals, ceratohyals, and thyrohyals); and bacular parameters (length of baculum, shape of base, and number, location, and size of lateral segments ). Detailed results of this study will be reported elsewhere. However, analy- sis of the data, including cluster analysis and principal components analysis, indicates the existence of an undescribed but well-differentiated subspecies of Microtus ochrogaster from central and southern Montana, Wyoming, and the contiguous comers of western South Dakota and Nebraska. Microtus ochrogaster similis, new subspecies Holotype.—Adult, female, skin and skull, No. 48183, Univ. Illinois Mus. Natur. Hist., from KULR-TV Tower, Billings, Yellowstone Co., Montana: collected June 15, 1973, by C. L. and C. W. Severinghaus and prepared by W. D. Severinghaus, original No. 1410. Range.—Central and southern Montana, central and northwestern \Wyo- ming, southwestern South Dakota, and northwestern Nebraska. Definition —A subspecies of Microtus ochrogaster characterized by: light brownish-black, hispid dorsal pelage, light to medium ochraceous ventral pelage; medium size, both externally and cranially; total length from 133.0 to 156.5 mm; hind foot from 19.5 to 20.6 mm; condylopalatal length from 17.3 to 18.1 mm; length of diastema from 7.85 to 8.20 mm, interorbital breadth from 3.78 to 4.03 mm; zygomatic breadth from 11.7 to 11.9 mm; condylobasilar length from 24.8 to 25.6 mm; palatilar length from 12.6 to 13.3 mm; prelambdoidal breadth from 9.03 to 9.21 mm; and depth of brain- case from 7.58 to 7.90 mm. Diagnosis —Microtus ochrogaster similis differs from M. ochrogaster hay- denii, the only subspecies with which it integrades, in being smaller 50 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Table 1. External and cranial measurements as population minima and maxima of the subgenus Pedomys. similis haydenii minor ochrogaster ohionensis ludovicianus Total 133.0 143.7 110.0 132.0 122.8 143.1 length 156.5 165.6 132.0 oles 149.0 Hind foot 19.5 19.3 15.0 IY 53 7a 18.2 length 20.6 PALS) ALO) 20.0 19.0 Condy!obasilar 24.8 24.3 19.8 SoD) QZIo 24.8 length VAC) 26.8 Dn Dom 24.2 Palatilar 12.6 1), 7 9.8 11.9 11.6 DA length SES 14.1 11.4 NBs) WD IL Condylopalatal 17.3 17.3 14.0 168 16.1 17.4 length ISI 19.2 15.4 18.0 AO Length 7.85 7.58 6.25 7.60 138 WO diastema 8.20 8.95 7.00 8.49 Cod Maxillary Gaal: 6.28 4.90 5.60 5.67 6.39 toothrow 6.43 6.83 5.48 6.32 5.07 Zy gomatic 14.7 14.2 10.8 13.9 MBL 7 14.4 breadth aS 15.9 12.4 aes 14.6 Interorbital 3.78 3.84 320 3.70 Be 3.83 breadth 4.03 4,31 3.60 4.19 4.02 Lambdoidal Loe 11.4 9.0 10.6 ILS} IIS) breadth WIL) 12.6 10.3 12.6 11.9 Prelambdoidal 9.03 9.37 7.80 8.72 8.83 9.14 breadth 9.21 10.00 8.40 9.85 9.20 Depth 7.58 UCAS 5.95 7.00 7.40 7.44 braincase 7.90 8.10 6.81 8.02 CeO Incisive 4.32 AL 3.40 4.10 4.13 4.70% foramina 4.63 5.03 4.14 4.77 AD * Includes taylori. + Only one population. (Table 1) in all external and cranial measurements, especially prelamb- doidal breadth and incisive foramina. Microtus ochrogaster similis differs from Microtus minor as follows: markedly larger (see Table 1); dorsal pelage a light brownish-black rather than dark brownish-black. Microtus ochrogaster similis differs from M. ochrogaster ochrogaster as follows: similar size except hind foot larger; hispid dorsal pelage light brownish- black rather than dark brownish-black. Microtus ochrogaster similis differs from M. ochrogaster taylori in the same way as does M. ochrogaster hay- denii. Microtus ochrogaster similis differs from M. ochrogaster ohionensis as follows: markedly larger; dorsal pelage a light brownish-black rather VOLUME 90, NUMBER 1 51 than dark brownish-black; ventral pelage light to medium ochraceous rather than white to gray. Microtus ochrogaster similis differs from M. ochrogaster ludovicianus as follows: larger; dorsal pelage a light brownish- black rather than dark brownish-black. Description—The holotype, an adult female, and six adult topotypes, four males and two females are, in millimeters, respectively: total length, 142, 151, 129, 135, 141, 152, 148; length of hind foot, 18, 20, 18, 19, 20, 21, 20; length of ear, 12, 13, 12, 14, 13, 14, 14; condylobasilar length, 25.4, 26.6, 23.1, 24.6, 25.6, 27.3, 26.0; length of diastema, 8.1, 8.6, 7.2, 7.7, 8.4, 8.9, 8.3; palatilarsength, 12.8, 13.1, 1.4, 12.2, 112°9, 113.5, 12:8: alveolar length of maxillary toothrow, 6.5, 6.7, 5.7, 6.3, 6.6, 6.9, 6.8; zygomatic breadth, 15.0, 15.9, 13.9, 14.6, 15.1, 16.6, 14.9; interorbital breadth, 3.7, 3.95, 3.9, 3.8, 3.5, 3.8; prelambdoidal breadth, 9.1, 9.2, 9.1, 9.15, 9.3, 9.4, 9.3: lambdoidal lmeadtinnmeleG. 12225 10. 11-6, °12:2, 1265 07; depth of brainease, 7.6, 7.9, (Poeeno Ono, 8.0; condylopalatal length, 17.7, 18.4, 16:1) 17.3, 17.8; 19:3, Se -leneth Or imcisive foramina, 5:2, 5.2, 4.7, 4.2, 4:6, 4:7, 5.1). Remarks.—Microtus ochrogaster similis differs appreciably from Microtus minor. Microtus o. similis integrades with M. o. haydenii in Custer and Fall River counties, South Dakota, Sioux County, Nebraska, eastern Mon- tana, and southern and eastern Wyoming. No such zone of intergradation, or sympatry, exists between M. o. similis and Microtus minor as approxi- mately 150 miles separate these two taxa. Microtus ochrogaster is a variable species but most of this can be ac- counted for by the recognition of subspecies. The external and cranial characters, excluding color, of Microtus ochrogaster similis resemble and are most similar to M. o. ochrogaster, not the contiguous M. o. haydenii. This is especially well shown by such parameters as condylobasilar length, zygomatic breadth, palatilar length, length of diastema, length of alveolus of the maxillary toothrow, and prelambdoidal breadth (Table 1). All of these are well within the range of variation of M. o. ochrogaster and only approach the range of variation of M. 0. haydenii. There is perhaps a slight clinal trend within M. ochrogaster from the small M. 0. ohionensis in the east, through M. o. ochrogaster, to the large M. o. haydenii in the west. It is somewhat surprising to find populations of the small sized M. o. similis in the western part of the range of M. ochrogaster. A principal components program, written by R. B. Selander, was used to analyze 154 pooled localities representing 1676 adult specimens. The first two principal components accounted for 93.46 per cent of the variation and the range in principal components scores for each taxon was deter- mined (Table 2). The range of scores for M. o. similis falls within the range of M. o. ochrogaster for the first and second principal components and only a small overlap with M. o. haydenii for the second principal com- ponent. 52 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Table 2. Principal component scores of the subgenus Pedomys. PC#I PC#2 similis 0.5606—0.6 152 0.3011—0.3156 haydenii 0.6183-0.7501 0.3121—0.3410* minor 0.0330—0.2877 0.2505—0.2841 ochrogaster 0.4459-0.6496 0.2973-0.3145 ohionensis 0.4430-0.5327 0.3214—0.3530 ludovicianus 0.5647 0.29697 * Includes taylori. * Only one population. A cluster analysis program written by F. J. Rohlf and modified by R. B. Selander was utilized to analyze 158 pooled localities (Fig. 1). The dendrogram produced by this program indicated that M. o. similis clusters with M. o. ochrogaster and M. o. ludovicianus as eight of 11 O.T.U.’s representing similis clustered with ochrogaster while the remaining three clustered with haydenii. The type specimen for this subspecies was collected by C. L. and C. W. Severinghaus in an old field at the base of a television tower south of the Yellowstone River at Billings, Yellowstone Co., Montana on June 15, 1973. The field had short grass and weeds with no noticeable runways. Three specimens, two females and one male, were taken on June 15 and five additional specimens, four males and one female on June 16. The subspecific epithet for M. o. similis reflects its similarity with M. o. ochrogaster. ochrogaster /UGOVICIONUS SITTUIIS OAIONENS/S hayden! taylor: Minor OF tO: Orr OD e204 OS: VOxe, BOR 0.0 Average Taxonomic Distance Fig. 1. Dendogram of the subgenus Pedomys. Note close relationship of similis to ochrogaster as compared to haydenii and minor. VOLUME 90, NUMBER 1 53 Specimens Examined The following abbreviations have been used: University of [Illinois Museum of Natural History (UIMNH), U.S. National Museum of Natural History (USNM), University of Michigan Museum of Zoology (UMMZ), University of Kansas (KU), Michigan State University (MSU), and Indiana State University (ISU). All collections listed are M. o. similis. MONTANA. Phillips Co.: 4 mi N James Kipp State Park, 3 (UIMNH). Garfield Co.: Piney Buttes, 10 (USNM); Jordan, 1 (UMMZ). Prairie Co.: petienommleny, 20) (USNM). Wheatland €o:; 4 mi SE Shawmut, 3 (UMMZ). Musselshell Co.: 16 mi S Roundup, 2 (USNM). Rosebud Co.: Ingomar, 3 (USNM); 28 mi NE Porcupine Creek, Hole in the Rock, 1 SUSNMGEeLame Deer, 2 (USNM). Custer Co:: Tongue River, 3 mi S Miles City, 17 (USNM ); Fort Custer [= Fort Keogh], 18 (USNM). Sweet Grass Co.: 13 mi ENE Melville, 2 (UMMZ); 11 mi ENE Melville, 8 (UMMZ). Stillwater Co.: Columbus, 6 (USNM); 10 mi NW Park City, 1 (USNM). Carbon Co.: 3 mi E Bear Creek, 4 (UMMZ). Yellowstone Co.: Lake Basin, 2 (USNM); Billings, 9 (ISU), 1 (USNM); KULR-TV Tower, Billings, 8 (UIMNH); Indian Creek, 8 mi SE Huntley, 4 (USNM). Big LlonmmCor 4 mi S Guster, 1 (KU); 15 misS ‘Custer, 7 (KU); Toluca, 6 (USNM); Crow Agency, 125 (USNM); Busby, 1 (KU); 20 mi N Wyo- ming, Little Bighorn River, 2 (USNM); 2 mi N Wyoming, Little Bighorn River, 1 (USNM); Valley of Little Bighorn River, 4 (USNM); Little Bighorn River, 1 (USNM). Powder River Co.: Powderville, 3 (KU), 3 (MSU); Powder River, Powderville, 7 (USNM); Moorhead, 12 (USNM): Powder River, Moorhead, 5 (USNM). No County: Crazy Mts., 2 (USNM). SOUMA DAKOTA. Custer Co.2 7 mi E faith, 1 (UIMNH). Fall River Co.: 9 mi S, 3 mi W Oelrichs, 2 (KU). WYOMING. Park Co.: 4 mi N Garland, 2 (KU). Big Horn Co.: Grey- bull; 9 (USNM); 8/10 mi S Greybull, 2 (KU); 1 mi S Greybull, 1 (KU). Washakie Co.: 3 mi E, 1 mi N Ten Sleep, 1 (KU). Sheridan Co.: Dayton, 1 (USNM); 3 mi WNW Monarch, 3 (KU); 4 mi NNE Banner, 4 (KU); 5 mi NE Clearmont, 12 (KU). Johnson Co.: 1 mi W, 8/10 mi S Buffalo, 5 (KU); 15% mi W, 1 mi S Buffalo, 1 (KU); 4% mi W, 1 mi S Buffalo, 1 (KU); 6% mi W, 2 mi S Buffalo, 3 (KU); % mi E Klendike [= Klondike], 1 (KU); 1 mi WSW Kaycee, 5 (KU). Campbell Co.: Ivy Creek, § mi W, 5 mi N Spotted Horse, 4 (KU). Crook Co.: 1% mi NW Sundance, 1 (KU); 15 mi ENE Sundance, 1 (KU); Sundance, 2 (USNM). Weston Co.: 1 mi N Black Hills, 1 (UMMZ); Newcastle, 1 (USNM); 23 mi SW Newcastle, 2 (KU). Fremont Co.: 12 mi N, 3 mi W Shoshoni, 11 (KU); 1% mi W, 2% mi S Casper, 3 (KU); 6 mi S, 2 mi W Casper, 1 (KU); 2 mi W, 7m S$ Casper, 1 (KU); Alcova, 1 (KU); Sun, 1 (USNM). Converse Co.: 4 mi W, 6 mi N Bill, 1 (KU); Beaver, 1 (USNM); 3 mi N, 5 mi E Orin, 9 (KU); 54 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Sand Creek, 1 (USNM). Niobrara Co.: 10 mi N Hat Creek, 9 (KU); 1 mi W tusk, 17 (KU); 2 mi'S, % mir E Lusk, 2 (KU). Carbon!@o;meass 4 (USNM ). NEBRASKA. Sioux Co.: 8 mi N Harrison, Warbonnet Twp., 4 (UMMZ); pvope, 8) (UR). ECU EM). Acknowledgments Funds to defray the cost of travel, equipment, supplies, and computer time were supplied by the University of Illinois, Museum of Natural His- tory, the Department of Ecology, Ethology, and Evolution, and a Biomedi- cal Sciences Support Grant. I would like to extend my appreciation to Dr. Donald F. Hoffmeister for his help and direction during the course study and the writing of this manuscript. Museum of Natural History, University of Illinois at Urbana-Champaign 61801. PROG? BIO“; SOC, WASEL 90(1), pp. 55-60 OPHIACANTHA CLYPEATA N. SP. FROM THE BERING SEA, WITH A REDESCRIPTION OF OPHIACANTHA RHACHOPHORA CLARK (ECHINODERMATA: OPHIUROIDEA ) Michael A. Kyte An examination of the holotype of Ophiacantha rhachophora Clark, 1911 (USNM 25630), from the Korea Strait revealed definite discrepancies between it and Clark's original description. Clark’s account consisted of a complete drawing and a very brief comparison of O. rhachophora to O. levispina Lyman, 1878, and O. adiaphora Clark, 1911. Further material from Clark’s original series of O. rhachophora was obtained in an effort to match Clark’s specimens to his published description. The additional specimens had been collected from the Bering Sea and although similar to O. rhachophora, were significantly different in several aspects. Because of these differences and the geographical separation of the 2 groups of specimens, the Bering Sea animals are described herein as a new species. Additional U.S. National Museum specimens of O. rhachophora trom near the type-locality, Albatross Sta. 4902 (32°30’50’N, 128°34’40”E ), were then studied. Most of these closely resembled the type of O. rhachophora. How- ever, 3 samples labeled O. rhachophora were reidentified as follows: USNM 26985, 3 specimens of O. adiaphora; USNM 27060, 4 specimens of O. adiaphora; and USNM 26605, 1 specimen of O. rhachophora, and 1 of O. adiaphora. Matsumoto (1917) published a brief description and figures of specimens that he had identified as O. rhachophora. His figures resembled Clark's type-specimen quite closely except for minor differences. Ophiacanthidae Ophiacantha rhachophora H. L. Clark, 1911 JeneRs, IE 2 Redescription—Disc diameter 5.0 mm, arm length approximately 20-30 mm (no intact arms). Aboral surface of disc thickly covered with minute thorny stumps. Radial shields covered, only distal tips bare aborally. Upper arm plates triangular, widely separated, without spines. Oral interbranchial spaces covered with thorny stumps like those on disc except immediately distal to oral shield where scales are visible. Genital slits large, conspicuous. Oral shield variable, triangular with small distal obtuse lobe to pentagonal. Adoral shields relatively massive, wider without than within, sometimes bearing 1 thorny papilla. Oral shield and first lateral arm segment separated by adoral shields. Each jaw with 6 oral papillae and 1 infradental papilla: distal oral papillae markedly thorny; more proximal papillae flatter and 56 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Figs. 1, 2. Ophiacantha rhachophora. 1, Aboral view; 2, Oral view. broader, sometimes bifid or trifid at tips. First oral arm plate small, pentag- onal, slightly wider than long, succeeding plates larger, triangular, wider than long, well separated. Lateral arm plates broad, meeting both orally and aborally, separating oral and aboral arm plates. Lateral plate bearing 6 to 8 arm spines. Spines approximate aborally on first 2 basal segments beyond the disc. Lower arm spines very thorny with definite thorns ex- tending to tip of spines. Serration becoming less extensive on upper spines and occurring only at bases on uppermost on segments beyond first basal. VOLUME 90, NUMBER 1 Ul ~l Upper spines on first segment thorny to tip. Second spine from top longest, about 2 to 3 joints long. Tentacle scale single, slender pointed, markedly thorny. Distribution Japanese outer sublittoral and upper bathyal, 115-923 m (Clark, 1911; Matsumoto, 1917). Discussion.—Because of the variability of this species and the confusion surrounding it, the above description applies to both Clark’s original holo- type, USNM 25630, and those of Clark’s original specimens that I have not reidentified as another species. The most representative of these are USNM 26704 (from Albatross station 5091), and USNM 26048 (from Albatross station 4893). The above description differs from Clark's (1911) original description and figures in several points. First, the arm spines on the most proximal arm segment approximate on the aboral side of the arm and the uppermost arm spines have thorny bases. Secondly, the genital slits are large and con- spicuous. Finally, the oral shields are definitely triangular, longer than wide with a lobe distally joining with the oral interbranchial space and separated from the first lateral arm plates by the aboral shields. Matsumoto’s (1917) description of O. rhachophora differed from that of the type-specimen in stating that the outermost oral papillae arise from the adoral shields and one scale-papilla occurs on the first oral arm plate, projecting inwards and vertically. However, this papilla is not figured and its placement is not clear. Matsumoto’s figures show that the highest arm spines are smooth at their bases and thorny at the tips. In the type-specimen the spines are smoothest at their tips and more thorny at the bases, except for those on the first aboral segment. These variations are not important enough to separate further specimens of this species. One of Clark's para- types, as discussed above, USNM 26704, possesses the thorny papilla arising from the adoral plate. Additional specimens labeled O. rhachophora trom 3 Albatross stations in the Bering Sea differ significantly from the type of O. rhachophora. These differences are both geographic and morphological and form the basis of a new species, O. clypeata. Ophiacantha clypeata, new species Figs. 3, 4 Etymology.—Clypeus (Latin), shield; referring to the shape of the aboral arm plates. Description.—Disc diameter 4-5 mm; arm length approximately 25-30 mm. Aboral surface of disc completely covered with thorny stumps; radial shields and all scales completely hidden. Upper arm plates vary from triangular with distal edge rounded, to shield-shaped, widely separated; first 2 or more basal upper arm plates bearing thorny stumps like those 58 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Figs. 3, 4. Ophiacantha clypeata. 3, Aboral view; 4, Oral view. covering disc. Oral interbranchial spaces more sparsely covered with stumps, scales partly visible especially near oral shields. Genital slits small, inconspicuous. Oral shields quadrangular, slightly wider than long. Adoral shields relatively massive, rectangular. Oral shield and first lateral arm plates not separated by adoral shields. Each jaw with 6 to 10 blunt oral papillae and 1 infradental papilla. All oral papillae slightly serrate, distal papillae slightly more so. First oral arm plate variable, of small to medium size, triangular to pentagonal. Succeeding plates larger, pentagonal, corners rounded, shallow notch sometimes in distal side. Lateral arm plates broad, VOLUME 90, NUMBER 1 59 meeting both orally and aborally, separating oral and aboral arm plates; each lateral plate bearing 6 arm spines. Lower arm spines very thorny. Serration decreases on ascending spines, top spine entirely smooth, longest 2%2-3 arm joints. Tentacle scale single, spiniform, slightly thorny at tip. Color, dried and in alcohol, brownish white. Distribution.—Bering Sea, collected at Albatross stations 4771, 4772, and 4775; “Bowers Bank”; upper bathyal, 629-1,068 m (Clark, 1911). Types.—Holotype: USNM 26241 from Albatross station 4771. Paratype: USNM 26698, from Albatross station 4772. Discussion.—Ophiacantha clypeata differs from O. rhachophora in having the distal tips of the radial shields concealed, the basal upper arm plates with thorny stumps, small and inconspicuous genital slits, oral shields quadrangular and wider than long, first lateral arm shields and oral shields contiguous, not separated by the aboral plates, arm spines on the first basal segment not meeting on the aboral side of the arm segment, and entirely smooth uppermost arm spines. While the oral papillae of O. rhachophora are markedly thorny, the 6 to 10 papillae of O. clypeata are only slightly serrate. A papilla does not arise from the adoral shield in O. clypeata. Geographical distribution supports the evidence from morphological differences. While O. clypeata occurs only in the Bering Sea, O. rhacho- phora is apparently restricted to Japanese waters and the Eastern Sea ( Clark, 1911; Matsumoto, 1917). Although the depth ranges of the 2 species overlap, O. rhachophora is found in somewhat shallower water than O. clypeata. This geographical separation and the morphological differences indicate clearly that Clark’s (1911) Bering Sea specimens of O. rhachophora should be separated as a distinct species. Acknowledgments I wish to thank especially Dr. David L. Pawson, Dr. James E. McCauley, and Dr. Andrew G. Carey, Jr., for their advice and encouragement during the preparation of this paper. Financial support was provided by the ERDA Contract E(45-1)-2227, Task Agreement No. 12. RLO 2227-T12-11. Literature Cited Clark, Hubert Lyman. 1911. North Pacific ophiurans in the collection of the United States National Museum. Bull. USNM 75:1—295. Matsumoto, H. 1917. A monograph of Japanese Ophiuroidea arranged according to a new classification. Jour. Coll. Sci., Tokyo 38(2):1—408. School of Oceanography, Oregon State University, Corvallis, Oregon 97331. Present address—Dames and Moore Environmental Sciences, Suite 500 Northgate Executive Center, 155 N.E. 100th Street, Seattle, WA 98125. PROC. BIOL. SOC. WASH. 90(1), pp. 60-65 A NEW SPECIES OF ELEUTHERODACTYLUS (ANUBA: LEPTODACTYLIDAE) FROM THE CORDILLERA ORIENTAL OF COLOMBIA William E. Duellman and John E. Simmons The high Andes of northern South America support a distinctive vege- tational formation, the paramo, characterized by a variety of low herbaceous plants, bunch grasses, cushion plants, some low woody bushes ( Baccharis, Compositae ), and the endemic composite Espeletia. Paramo occurs prin- cipally at elevations above 3,000 m, although in some areas it descends to about 2,500 m. The paramos receive abundant precipitation throughout the year; at higher elevations part of the precipitation falls as snow. Paramo communities are most extensive in Colombia and Ecuador and also occur in the Merida Andes of Venezuela. In contrast with adjacent lower montane habitats, the herpetofauna of the paramos is depauperate, consisting mostly of anurans of the genera Atelopus, Colostethus, Eleutherodactylus, Gastrotheca, and Phrynopus, al- though species of salamanders ( Bolitoglossa), teiid lizards (Anadia), and one other frog ( Hyla labialis) do occur in the paramos in Colombia. Among the herpetological inhabitants of the Colombian paramos, the genus Eleu- therodactylus is the most diverse in numbers of species. Frogs of this genus provide excellent subjects for the study of patterns of speciation in the paramos, which are distinctive habitat islands scattered through the three ranges of the Andes in Colombia. The frogs under consideration here are from the eastern range, the Cordillera Oriental. Upon studying large series of eleutherodactyline frogs obtained in the course of a survey of herpetological communities in the paramos in 1974, we discovered that one species well represented in our collections was unnamed. We take this opportunity to name this species for our friend and colleague, John D. Lynch of the University of Nebraska, in recognition of his devotion to the systematics of Eleutherodactylus. Eleutherodactylus lynchi, new species iG Holotype —KU 168139, an adult female, from Vado Hondo (05°26’N, 72°44’W ), 2660 m, Departamento Boyaca, Colombia, one of a series obtained on 28 August 1974 by William E. Duellman, Dana K. Duellman, John E. Simmons, and Linda Trueb. Paratopotypes—FMNH 202672-77, KU 168140-98, 168204-301, RMNH 17815(6), MLS 469-472, INDERENA (4), same collectors, 28 August— 2 September 1974. VOLUME 90, NUMBER lI 61 Fig. 1. Variation in color pattern in Eleutherodactylus lynchi: a, KU 168139 (holo- type), 2, 29.9 mm SVL; b, KU 168140, 9, 28.1 mm; c, KU 168141, 9, 31.4 mm; Gee G6S814250% 30:3 mm. Diagnosis —A member of the Eleutherodactylus unistrigatus group, char- acterized by: (1) skin on dorsum shagreened with scattered low granules, on belly coarsely granular; (2) snout subacuminate in dorsal view and round in profile, barely protruding beyond margin of lip; (3) width of eyelid about two-thirds of interorbital distance; (4) cranial ridges absent; (5) tympanum round with a moderately heavy supratympanic fold; (6) first finger slightly shorter than second; (7) fingers bearing lateral fringes: (8) terminal digital discs expanded, truncate; width of disc on third finger equal to diameter of tympanum; (9) palmar tubercle moderately small, elliptical; (10) forearm bearing ventrolateral row of low tubercles; (11) tarsus having scattered low tubercles, inner tarsal fold curved on distal two-fifths of tarsus; (12) inner metatarsal tubercle large, ovoid; outer metatarsal tubercle small, conical; (13) toes bearing lateral fringes and expanded terminal discs; outer toes webbed basally; (14) prevomerine odontophores small, behind level of choanae; (15) vocal slits and median subgular vocal sac present; (16) dorsum brown or tan with or without darker or paler markings; venter cream with or without brown flecks on chin and chest; upper lips and posterior surfaces of thighs plain brown. Eleutherodactylus lynchi differs from all other eleutherodactylines known from the high elevations of the Cordillera Oriental by having small pre- vomerine odontophores covered by buccal lining; in the other species the odontophores are distinct and larger. Furthermore, EF. /ynchi has expanded 62 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON terminal discs on the digits, which distinguish it from E. nicefori and Phrynopus nanus, both of which have narrow digital tips; the latter also lacks a tympanum. From the remaining species of Eleutherodactylus in the Cordillera Oriental, E. lynchi differs by having a curved tarsal fold that is continuous with the inner metatarsal tubercle. In E. bogotensis and E. verte- bralis the tarsal fold is short, straight and separated from the inner metatarsal tubercle; the tarsal fold is absent in E. affinis and E. elegans. Description—Body robust; head slightly wider than long; snout sub- acuminate in dorsal view, round in profile; snout barely protruding beyond margin of lip; top of head flat; width of eyelid about two-thirds of inter- orbital distance; canthus rounded; loreal region barely concave; nostrils slightly protuberant dorsolaterally; internarial area flat; lips not flared. Tympanum round, distinct, separated from eye by distance equal to diameter of tympanum, bordered above by moderately heavy supratympanic fold. Forearm moderately robust, bearing ventrolaterally a row of low tubercles; first finger slightly shorter than second; fingers having lateral fringes ex- tending to discs, which are nearly twice as wide as digit; subarticular tubercles large, subconical; palmar tubercle moderately small, elliptical; pollical tubercle elongate. Hind limbs short, robust; row of low tubercles on ventrolateral edge of tarsus; inner tarsal fold distinct, curved, on distal two-fifths of tarsus; inner metatarsal tubercle moderately large, ovoid; outer metatarsal tubercle subconical; toes slender, bearing lateral fringes; discs expanded, slightly smaller than those on fingers; subarticular tubercles small, subconical. Skin on dorsum shagreened with scattered small granules; mixture of large and small granules on flanks; two or three large round tubercles posteroventral to tympanum; skin on belly and ventral surfaces of thighs granular; skin elsewhere smooth. Anal opening a puckered aperture at midlevel of thighs, bordered laterally and ventrally by small tubercles. Pre- vomerine odontophores small, inconspicuous, covered by buccal lining, inclined posterolaterally behind the posterior borders of small, ovoid choanae. Vocal slits and single median subgular vocal sac present. Coloration in preservative-—The dorsum varies from tan to dark brown. Four color morphs can be discerned in the 240 known specimens: (1) Dorsum tan to brown with small irregular darker brown spots (62%); (2) Same as preceding except having a pale spot on the snout and another in the scapular area (13%); (3) Dorsum pale tan with or without brown streaks; flanks dark brown (20%); (4) Dorsum dark brown; flanks tan (5%). A narrow cream middorsal line is present in 9.6% of the specimens. The venter is cream; the posterior surfaces of the thighs are uniform brown. Coloration in life —The dorsum is dark brown with or without tan spots or cream middorsal line, or olive-green, tan, or orange-red with dark brown flanks. The posterior surtaces of the thighs are orange-brown; the belly is §3 VOLUME 90, NUMBER I GAG) SOE, S10 += 88& GLO + 9S 810 = GEV 1cO + 807 6V | + 9GE VIS —SLe cOr 996° ILE Gee" ILy 907 ISy -1L&- V 9E—-E'66 LI 6 Cc6E° ese’ OSe’ Cer vIV ccc Ley Loe S6e°-T Le" 98 -TLe" CVV —8CV vey OOF G6 LG—0 IG V r TOOEERS VOr 6LE coe" VYV SGV G1t LEV —68t" 66E"—89E SLE Soe" Soy —1&y 9SV G6" G £E—-9 6G € 6) ceO + GhY 800° + SLE oL0) == LOS 0cO’ + SIV SOS Salita Sol + 6G SSP -P6e S8E—C9e" OSE —9Se" ISP'—O08t" StPV -88o" GLG-& EG 6 y spite L90) == 1GV O10 + 88 LO) SEs OLS VIO + 6¢6F LIO’ == 907 OG I + 986 £99066 CIV VLE GGV OSE 8SV' 66" try OLE: 8 1E-L 96 SG 6 GSO’ = PHY C10) == OLE 600° + 99€" 160 += 1VV 610 + CIV IST = SVG 989° —S8e" COV 8h O08 —6Se" cLVy —6O0V 9SV C8" 6 L6-9 GS LI 7 OY PE Gs CA AN SUSE TAS TAS TAS IAS TAS N 2S AWeooy /ummueduts J, /YIPIA prop /YAsusy peroy{ /YSusy] 1004 /Y{s8ug] eIqiL, ‘UOI}WIAOp piepuejs [ = UPROUL ST OUT] PUOOS ‘UOHRIIVA FO OBUBI SI OUT] FSI “Wy IUliy snphzopposayznayy fo suonsodo1d pue (TAS) YIsuey, yUaA—jnoUS TE SISMeIL 64 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON gray, and the vocal sac is dull yellow. The iris is dull bronze with a median horizontal brown streak. Measurements.—The holotype, an adult female, has the following di- mensions (in mm): snout—vent length 29.9, tibia length 12.4, foot length 13.7, head length 11.4, head width 12.1, diameter of eye 3.7, diameter of tympanum 1.3. Measurements and proportions of series from three localities show little variation (Table 1). Distribution —The type-locality is in the southeastern part of the Paramo de Vigajual; Eleutherodactylus lynchi was found at 2,460 m, 6 km S of Vado Hondo and at 3,150 m, 20.5 km NW of Vado Hondo, both in the Paramo de Vigajual. The species was also collected in the Paramo de la Rusia (3,340 m), 19 km NW of Duitama and in paramos 22 km SSW of Susacon (3,000 m) and 33 km NE of Belén (2,880 m). All localities are in the Cordillera Oriental in the Departamento de Boyaca. Remarks.—Eleutherodactylus lynchi belongs to Group II of the genus as defined by Cochran and Goin (1970) and to the large Eleutherodactylus unistrigatus group as defined by Lynch (1976). This group is most diverse in the Andes of Colombia and Ecuador. In the Cordillera Oriental this group is also represented by (1) E. nicefori in the Paramo del Almorzadero to the north of the range of E. lynchi; (2) E. affinis, bogotensis, and elegans in the paramos in the vicinity of Bogota to the south of the range of E. lynchi; (3) E. vertebralis widespread in paramos, subparamos, and upper montane forests in the three Colombian cordilleras southward into Ecuador. Only E. vertebralis occurs sympatrically with E. lynchi (Paramo de la Rusia). Vado Hondo is a small (four houses) village on the road east of Sogamosa. Our study site was just north of the village, across the Rio Cusiana. Along the northern and western sides of the river are steep slopes with granitic talus. The vegetation consists of grasses (partially grazed by cattle and goats), many lichens and mosses, and scattered clumps of Espeletia and Baccharis. In some areas mosses formed about 70% of the ground cover. During the five days that we spent at Vado Hondo the temperature varied from nocturnal lows of 4.0-7.0 (x = 5.4)°C to diurnal highs of 12.0-19.5 (x = 13.7)°C. Rain fell daily, accumulating to 6-16 (x = 12.2) mm daily. Although some specimens of E. lynchi were on the valley floor, most were under rocks on, or at the base of, the south and east facing slopes. All were found by day. Six females having snout—vent lengths of 26.6-33.3 (x = 30.1) mm contained 6-16 (x = 10.8) unpigmented ovarian eggs 1.2— 1.8 (x = 1.5) mm in diameter. The testes are white in 75% of the adult males and heavily spotted with black or entirely black in the others. At Vado Hondo and Paramo de la Rusia E. lynchi is a member of a herpetofaunal community containing four other species of frogs (Atelopus ebenoides, Colostethus subpunctatus, Hyla labialis, Phrynopus nanus) and one salamander (Bolitoglossa sp.). Eleutherodactylus vertebralis is also present at Paramo de la Rusia. VOLUME 90, NUMBER 1 65 Specimens examined.—Abbreviations: Field Museum of Natural History (FMNH), Instituto Desarrollo de los Recursos Naturales Renovables, Bogota (INDERENA), University of Kansas Museum of Natural History (KU), Museo de la Salle, Bogota (MLS), Rijksmuseum van Natuurlijke Historie (RMNH), University of Michigan Museum of Zoology (UMMZ). All specimens are from Colombia. E. affinis: Cundinamarca: Barranquilla, UMMZ 57495. E. bogotensis: Cundinamarca: 11 km E Bogota (road to Choachi), KU 124693-739; Cerro Monserrate, Bogota, KU 110383-407, 124688-92; El Verjon, KU 150708-09. E. elegans: Cundinamarca: 11 km E Bogota, KU 124740-41; 15 km E Bogota, KU 132630. E. lynchi: Boyaca: 33 km NE Belén, KU 168348-60; Paramo de la Rusia, 19 km NW of Duitama, KU 168361-63; 22 km SSW Susacoén, KU 168315-47; Vado Hondo, FMNH 202672-77, INDERENA (4), KU 168139-98, 168204- 301, MLS 469-472, RMNH 17815(6); 6 km S Vado Hondo, KU 168302-14; 20.5 km NW Vado Hondo, KU 168364-83. E. nicefori: Norte de Santander: 18.5 km S Chitaga, KU 168445-80; 32 km S Chitagé, FMNH 202678-83, KU 168481-516, RMNH 17811(4). San- tander: Chitaga: 150710-13; Paramo del Almorzadero, KU 150714-24. E. vertebralis: Antioquia: Sonson, KU 150733. Boyaca: Paramo de la Rusia, 19 km NW Duitama, KU 168625. Narino: N shore Lago de la Cocha, KU 168643; La Victoria, KU 140308-09. 8 km NE Pasto, KU 168640; 12 km E Pasto, KU 168641-42. Norte de Santander: 18.5 km S Chitaga, KU 168626-30; 32 km S Chitaga, KU 168631-39. Santander: 35 km ENE Bucaramanga, KU 132722-23; Paramo del Almorzadero, KU 150731-32; Presidente, KU 150727-30. Acknowledgments Field studies in the Andes were supported by a grant (No. 1304) from the National Geographic Society. We are grateful to our field companions, Dana K. Duellman and Linda Trueb, for their efforts in obtaining specimens and data, and to John D. Lynch for his critical review of the manuscript. Collecting permits were generously provided by Jorge Hernandez of the Instituto Desarrollo de los Recursos Naturales Renovables, Bogota. Literature Cited Cochran, D. M., and C. J. Goin. 1970. Frogs of Colombia. Bull. U.S. Nat. Mus 288:xii + 1-655. Lynch, J. D. 1976. The species groups of the South American frogs of the genus Eleutherodactylus (Leptodactylidae). Occas. Papers Mus. Nat. Hist. Unis Kansas 61: 1-24. Museum of Natural History, The University of Kansas, Lawrence, Kansas 66045. PROC. BIOL. SOC. WASH. 90(1), pp. 66-88 SPIONIDAE (ANNELIDA: POLYCHAETA) FROM SAN FRANCISCO BAY, CALIFORNIA: A REVISED LIST WITH NOMENCLATURAL CHANGES, NEW RECORDS, AND COMMENTS ON RELATED SPECIES FROM THE NORTHEASTERN PACIFIC OCEAN William J. Light This paper is part of the San Francisco Bay Project being carried out under the auspices of the Department of Invertebrate Zoology, California Academy of Sciences. The ultimate aim of this effort is to cover all marine and estuarine invertebrates of the intertidal and subtidal benthos of the San Francisco Bay system, and to produce a series of identification manuals covering this fauna (see Light, 1976). The polychaetous annelids are being investigated by means of earlier records, examination of previously recorded species where possible, and studies of additional material col- lected from various sources. This study was supported by a grant from the National Science Foundation (BMS 74-23523). I am indebted to Dr. Kristian Fauchald, Allan Hancock Foundation (AHF), University of Southern California, for the generous loan of the valuable collection upon which Hartman (1954) based her discussion of San Francisco Bay polychaetes. I am also grateful to Drs. Marian H. Petti- bone, National Museum of Natural History (USNM), Ernst Kirsteuer, American Museum of Natural History (AMNH), James A. Blake, Pacific Marine Station Museum (PMSM), University of the Pacific, Dillon Beach, California, and C. E. O'Riordan, National Museum of Ireland (NMI), for loans of type-specimens and other important material. Dr. Frederic H. Nichols, U.S. Geological Survey, Menlo Park, California, made available an extensive collection of San Francisco Bay polychaetes. The U.S. Army Corps of Engineers and the Bureau of Reclamation also provided major collections from the Bay and Bay-Delta region, as well as from the Delta- Mendota Canal. These collections have been accessioned into the Depart- ment of Invertebrate Zoology, California Academy of Sciences (CAS). Dr. James A. Blake shared his thoughts on many of these issues and generously allowed me to use some of his unpublished data. Both he and Dr. Keith H. Woodwick, California State University at Fresno, kindly reviewed the manuscript. Dr. Welton L. Lee, California Academy of Sciences, helped to make the prose more readable. The polychaete family Spionidae is well represented in San Francisco Bay, both in numbers of species, as well as numbers of individuals. More detailed information will be presented in an identification guide to the Spionidae soon to be published (Light, in preparation), the first of the VOLUME 90, NUMBER I 67 previously mentioned series of such manuals. The present paper deals with some of the systematic problems encountered, revisions of some of the names, new distributional records and comments on the species found in San Francisco Bay. The 19 species of Spionidae known to occur in San Francisco Bay are listed below, of which 10 are newly recorded. Three additional, closely related spionid species from the northeastern Pacific are added for comparison. The following Spionidae are covered in this report: B(N): E E ooo cose oreo ooo“ e 4 Z | B(N) B(N) B(N) B(N) B(N) B B(N) B Boccardia ligerica Ferronnicre, 1898:109.—Blake & Woodwick, 1971: Previously reported from San Francisco Bay Newly recorded for San Francisco Bay Boccardia ligerica Ferronnicre Boccardia proboscidea Hartman Polydora brachycephala Hartman Polydora ligni Webster Polydora quadrilobata Jacobi Polydora socialis (Schmarda ) Pseudopolydora kempi (Southern ) Pseudopolydora paucibranchiata (Okuda ) Pygospio californica Hartman Pygospio elegans Claparede Scolelepis squamata (Miller) (as Nerine cirratulus Nerinides acuta) Scolelepis tridentata (Southern) Scolelepis alaskensis (Treadwell) Dispio uncinata Hartman Spiophanes berkeleyorum Pettibone Spiophanes kroeyeri Grube Spiophanes bombyx (Claparede ) Prionospio (Prionospio) steenstrupi Malmgren Prionospio (Apoprionospio) pygmaea Hartman Prionospio (Minuspio) cirrifera Wiren Paraprionospio pinnata (Ehlers ) Streblospio benedicti Webster Boccardia ligerica Ferronnicre fig. 1 [synonymy]. Polydora redeki Horst, 1920:111.—Hartman, 1941:290; 1969:97.—Reish & Winter, 1954:113. Boccardia nr. uncata: Hartman, 1954:9 [Not Berkeley, 1927]. Polydora uncata: Filice, 1958:170, 190 [Not Boccardia uncata Berkeley, 1927]. and Q9 IL, 68 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Material—California: San Francisco Bay: San Pablo Channel, several specimens (AHF n10173-75; as Boccardia nr. uncata by Hartman). Off Benicia, Carquinez Strait, 1 specimen (CAS 000715). Suisun Bay, 1971, 40 specimens (CAS 000724). Honker Bay, August 1973, 31 specimens (CAS 000716-18). Delta-Mendota Canal, milepost 10.62, in fresh water, Novem- ber 1973, 1 specimen (CAS 000717). Remarks.—New record for San Francisco Bay and central California. Boccardia uncata Berkeley was referred to B. hamata (Webster) by Blake (1966:177). Although B. hamata is commonly collected outside San Francisco Bay in such localities as Bolinas Lagoon, Marin Co. (Light, un- published data), it is not known to occur inside the Golden Gate Bridge. The specimens previously reported from San Francisco Bay by Hartman (as B. nr. uncata) and Filice (as P. wncata) are herein referred to Boccardia ligerica. The 2 species are distinguished as follows: B. ligerica bears branchiae on setigers 2, 3, 7 and subsequent segments, and the pygidium consists of an undivided ventral plate bearing 2 anal cirri (see Blake & Woodwick, 1971, fig. 1i). B. hamata has branchiae on setigers 2, 3, 6 and subsequent segments, and the pygidium comprises 2 ventral lappets, each with a small terminal papilla (see Blake, 1966, fig. 7). B. ligerica has been reported from southern California as B. redeki by Hartman (1941, 1969) and by Reish & Winter (1954), although these records have not been con- firmed (see Blake & Woodwick, 1971:34). The species is otherwise known from brackish waters and mud flats of western Europe. Boccardia proboscidea Hartman Boccardia proboscidea Hartman, 1940:383; 1954:9; 1969:95. The occurrence of this species in San Francisco Bay is not clear. It was listed by Hartman (1954, 1969) as occurring in the Point Richmond area on the authority of S. F. Light (1941:63). However, Light merely lists B. proboscidea as occurring in San Francisco Bay [as indicated by the letter “B” on p. 63]. His key to the Annelida from Point Richmond on p. 181 included only the genus Boccardia with no reference to a given species. With the exception of this uncertain record, this species has not been re- ported from San Francisco Bay. Polydora brachycephala Hartman Fig. la-c Polydora brachycephala Hartman, 1936:48; 1969:129.—Filice, 1958:190.— Reish, 1959a:38; 1964:86-92.—Blake, 1971:11 [synonymy]; 1975:214. Polydora caulleryi: Pettibone, 1954:280 (part).—Banse, et al., 1968:540 (part ).—Blake, 1971:11 (part). [Not Mesnil, 1897. ] VOLUME 90, NUMBER 1 69 b C (ue eet oh aig aL (O72 Fig. 1. Polydora brachycephala Hartman. Modified major spines from setiger 5 of aberrant juvenile specimen (CAS 000723), showing various stages of growth from oldest (a) to youngest (c). Scale in mm. Material—Calitornia: Polydora brachycephala Hartman, Drakes Estero, Marin Co., 1933, 2 syntypes plus fragments (CAS Type Series 680). San Francisco Bay: Hunter's Point, Potrero, Foster City, Oakland Airport, Oak- land Inner Harbor, Alcatraz Island, San Pablo Bay, Tiburon, 1973-1974, many specimens (CAS 000719-21, 000726-29, 000734-37, 000739-50 ). South San Francisco Bay, September 1973, 1 aberrant juvenile specimen (CAS 000723 ). Remarks.—Polydora brachycephala was placed into synonymy with P. caulleryi by Pettibone (1954:280). This view was accepted by Banse, Hob- son & Nichols (1968) and Blake (1971). However, P. brachycephala has been retained as a valid species by Reish (1959a, 1964), Hartman (1969) and Blake (1975). As a result of recent investigations, J. A. Blake (per- sonal communication) has noted substantial differences in larval develop- ment between examples of P. caulleryi from the east coast of North America and P. brachycephala from the west coast. Because other Polydora species with distributions on both the east and west coasts do not show such differences, he now prefers to retain P. brachycephala as a species distinct from P. caulleryi until further life history information is available. This view is adopted here, and the name P. brachycephala is provisionally retained for the central California population. This species was_pre- viously reported from San Francisco Bay by Hartman (1936, 1954) and Filice (1958). An aberrant juvenile specimen (CAS 000723) was found which exhibited 2 kinds of modified spines on setiger 5: the more usual caulleryi- brachycephala type of spine (Fig. la) blends into a more slender falciger with a long distal tip (Fig. lb-c); both types bear a subdistal, sparse brushy tip. The companion setae are similarly fimbriated. The specimen is unique 70 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON in bearing hooded hooks in the neuropodia from setiger 6 on one side, and irom setiger 7 on the other. The first occurrence of the hooded hooks is usually quite constant within a species. Polydora ligni Webster Polydora ligni Webster, 1879b:119.—Hartman, 1941:310; 1954:10; 1969: 137.—Graham & Gay, 1945:379.—Filice, 1958:167, 190.—Jones, 1961: 266, 268—Blake, 1971:5 [synonymy]; 1975:210.—Foster, 1971:22 [synonymy ]. Polydora amarincola Hartman, 1936:46. Polydora ligna [sic]: Filice, 1954:16. Material_—California: San Pablo Bay, Point Wilson, July 1951, sta. 16, coll. F. P. Filice, several specimens (AHF n10397). Off Standard Oil Refinery, Richmond, 1969, several specimens (CAS 000730, 000732-33 ). South San Francisco Bay, Redwood City Harbor, Hunter’s Point, Oakland Inner Harbor, 1973-1974, many specimens (CAS 000751-69 ). Remarks.—This species was first recorded from the east side of San Francisco Bay and Lake Merritt, Oakland, California by Hartman (1936) as a new species, P. amarincola. It was subsequently reported as P. ligni from San Francisco Bay and Lake Merritt by Hartman (1941), from Oak- land estuary by Graham & Gay (1945), from the Castro Creek region of San Pablo Bay by Filice (1954), from Carquinez Strait by Filice (1958) and off Point Richmond by Jones (1961). Polydora quadrilobata Jacobi Polydora quadrilobata Jacobi, 1883:31, pls. 1 & 2 (part).—Jones, 1961: 288.—Hartman, 1961:100; 1969:145.—Blake, 1971:13 [synonymy]. Material—British Columbia: Vancouver, Stanley Park (Lumberman’s Arch), 49°18’12”N, 123°07’33’W, from exposed mudflats, March & April 1969, W. J. Light, coll., several specimens (CAS 000728-29). Remarks.—This species was collected from San Francisco Bay only once by Jones (1961) and was represented by two individuals taken with an Eckman grab of Point Richmond in November 1955. According to Dr. M. L. Jones (personal communication), these specimens have been lost. This form has otherwise been reported from the eastern Pacific Ocean off south- ern California by Hartman (1961, 1969). Specimens were collected in vast numbers in British Columbia in the spring of 1969. This population exhibited explosive growth with a preponderance of juveniles at first, and with an increasing proportion of adults which persisted in huge numbers for several weeks, only to disappear as quickly as they appeared. The ani- mals constructed interconnecting mud galleries to a depth of several inches in the mud which terminated in erect tubes projecting above the substratum. VOLUME 90, NUMBER 1 he Polydora socialis (Schmarda ) Leucodore socialis Schmarda, 1861:64, pl. 64 figs. a—c, pl. 26: fig. 209. Polydora socialis plena Berkeley & Berkeley, 1936:468. Fide Blake, 1971:20. Polydora socialis: Hartman, 1941:310; 1969:147.—Blake, 1971:20, fig. 13 [synonymy]; 1975:215. Polydora plena: Foster, 1971:24, figs. 22-29. Material——California: San Francisco Bay, Hunters Point, December 1973, 1 specimen (CAS 000722). Hunter's Point, 1973-1974, 21 specimens (CAS 000770-78 ). Off Oakland & San Francisco airports, January-February 1973, 11 specimens (CAS 000779-82 ). Remarks.—New record for San Francisco Bay. Pseudopolydora kempi (Southern ) Polydora (Carazzia) kempi Southern, 1921:636, fig. 20a-j. Polydora (Carazzia) kempi var.: Okuda, 1937:233, figs. 13, 14. Neopygospio laminifera Berkeley & Berkeley, 1954:462. [See also Banse, IG R22ANS). | Pseudopolydora kempi japonica Imajima & Hartman, 1964:287. Pseudopolydora kempi californica Light, 1969:542. Pseudopolydora kempi: Hartman, 1969:167.—Blake, 1975:215.—Blake & Woodwick, 1975:126—Chapman & Dorman, 1975:107. Polydora (Pseudopolydora) kempi japonica: Banse, 1972:219. Material.—British Columbia: Nanaimo, Rathtreuer Beach, July 1951, 3 paratypes of Neopygospio laminifera (USNM 32716). California: Bolinas Lagoon, Marin Co., August 1967, holotype and 2 para- types of Pseudopolydora kempi californica (CAS Type-Series 377-379 ). San Francisco Bay, San Pablo Bay (Oleum), Hunter's Point, 1972, 3 speci- mens (CAS 000788-90). South San Francisco Bay, Redwood City Harbor, Oakland Inner Harbor, 1973-1974, many specimens (CAS 000791-99, 001881-88 ). Remarks.—Although this species is well known in central California, it has only recently been reported from San Francisco Bay by Chapman & Dorman (1975) and Blake & Woodwick (1975). Pseudopolydora paucibranchiata (Okuda) Polydora (Carazzia) paucibranchiata Okuda, 1937:231, figs. 11, 12.—Reish, 1954:255; 1955:1171; 1959a:38, 47; 1959b:S86—-S7.—Barnard, 1958: 167. Polydora (Carassia) [sic] paucibranchiata: Reish, 1954:224. Polydora paucibranchiata: Reish, 1961a:266; 1963:25. Pfolydora]. (C.) paucibranchiata: Reish, 1961b:S6. 72 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Pseudopolydora paucibranchiata: Imajima & Hartman, 1964:288.—Blake, 1975:215.—Blake & Woodwick, 1975:110. Pseudopolydora cf. paucibranchiata: Read, 1975:414, fig. 7. Material.—California: South San Francisco Bay, September & December 1974, Hunter’s Point & Oakland Inner Harbor, 1973-1974, numerous speci- mens (CAS 001889-1904 ). Remarks.—New record for San Francisco Bay. This species is well known in southern California and Tomales Bay, Marin Co. It is otherwise known from Japan [type-locality] and questionably from New Zealand (Read, Oia) Pygospio californica Hartman Pygospio californica Hartman, 1936:50, figs. 17-19; 1944b:260; 1954:10; 1969:169, figs. 1-3.—Blake, 1975:215. Material.—California: Moss Beach, San Mateo Co., 1933, 9 paratypes plus fragments (AHF Type 0612). Del Monte Beach, Monterey Co., May 1972, many specimens (CAS 000710). Remarks.—This species was reported once from San Francisco Bay off Point Richmond in a few feet of water by Hartman (1954). An examina- tion of the paratypes clearly shows this form to be distinct from P. elegans Claparede. P. californica bears an anteriorly tapering, bluntly conical and bulbous prostomium projecting well forward of the peristomium, a com- pletely eversible, saclike proboscis with distinct, diffuse brown spots, the branchiae are first present from setiger 17-21, and bidentate hooded hooks first appear in the neuropodia from setiger 23-25. P. elegans exhibits an anteriorly expanded prostomium, often with lateral swellings, which is never tapering and which may be slightly incised; the proboscis is only partially eversible, conical (never saclike) and without pigment spots; the branchiae are first present from setiger 11-13; and both bidentate and spoon-shaped hooded hooks are first present in the neuropodia from setiger 8-9 [see below]. Pygospio californica has otherwise been reported only from San Mateo and Marin counties, California. The range is here extended to Monterey Co. Pygospio elegans Claparede Pygospio elegans Claparede, 1863:37.—Soderstrom, 1920:267.—Berkeley & Berkeley, 1954:462.—Foster, 1971:29 [synonymy].—Hobson, 1974:71.— Blake, 1975:215.—Blake & Woodwick, 1975:124. Material_—California: San Francisco Bay, near Redwood City, 1 speci- men (CAS 000700). Marin Co., Dillon Beach, Pacific Marine Station over- flow pond, 25 June 1976, many specimens (CAS 002038). ——— VOLUME 90, NUMBER 1 73 Remarks.—New record for San Francisco Bay. This species has only recently been recorded from the eastern Pacific Ocean off Washington and British Columbia by Hobson (1974). Blake (1975) also mentions this form in his discussion of central California intertidal polychaetes, and Foster (1971:32) lists California under the distribution of this species. It is quite abundant in Tomales Bay, Marin Co. (Blake & Woodwick, 1975) and Bodega Harbor, Sonoma Co., California (J. A. Blake, personal communica- tion ). Soderstrom (1920:75-77, figs. 67-72) commented at length upon the peculiar spoonlike crotchets (“loffelformige Borsten” ) which he observed in the neuropodia of setigers 8-14 of mature P. elegans. He concluded that the development of these spoonlike forms was a function of age, and that they derived from a transformation of the more normal bidentate crotchets as the worm matured. He noted that the younger the individual, the fewer were the number of anterior crotchet-bearing neuropodia exhibiting these unusual setae. These spoonlike crotchets were subsequently reported in a single specimen from Nova Scotia by E. & C. Berkeley (1954:462, fig. 5): apparently they have not been observed since, and Foster (1971) does not mention them in her detailed account of this species. It is noteworthy, then, that all specimens available to me from central California exhibit spoonlike crotchets in setigers 8 through 10 or 12 precisely like those figured by Soderstr6m and the Berkeleys. Scolelepis squamata (Miller ) Figs. 2a-f, 3a—b Lumbricus squamatus Miller, 1806:39. Spio acuta Treadwell, 1914:199. Nerine minuta Treadwell, 1939:5. Nerinides agilis: Hartman, 1944a:340; 1956:291—Dean & Hatfield, 1963: 163. Nerine cirratulus: Berkeley & Berkeley, 1952:27.—Hartman, 1954: 10. Nerinides acuta: Hartman, 1954:10; 1969:119, figs. 14.—Filice, 1958: 1$9.— Jones, 1961:288, 314, 317.—Blake, 1975:214. Scolelepis squamata: Pettibone, 1963:92 [synonymy |.—Foster, 1971:60, figs. 118-131 [synonymy ]. Scolelepis squamatus: Blake, 1975:215. Material—Western Europe: Italy, Bay of Naples, 1 specimen (USNM 5135). Netherlands, near Dan Helder, 1953, 4 specimens (USNM 34211). East coast of North America: Hampton Beach, New Hampshire, 1963, 5 specimens (USNM 34218). Gulf of Mexico: Sarasota, Florida, 1960, 3 specimens (USNM 43036), Port Aransas, Texas, holotype of Nerine minuta (AMNH 2566). 74 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Th i si) .03 e f [| {OS Fig. 2. Scolelepis squamata (Miller), neuropodial hooded hooks. a, Tridentate hook in right lateral view; b, Bidentate hook in right lateral view; c, Unworn bidentate hook in frontal view; d, Tridentate hook in postero-lateral aspect with hood removed; e, Wom hook in frontal view; f, Worn hook in left lateral view. Scale in mm. British Columbia: Long Bay, August 1931, 1 specimen (USNM 40792; as Nerine cirratulus by E. & C. Berkeley ). California: San Diego, paratype of Spio acuta (AMNH 764). San Francisco Bay, lower section of bay, 9-26 m, Albatross sta. D5724-27, 5729-30, March 1912, and outside Golden Gate, 73 m, Albatross sta. D5787, October 1912, several specimens (AHF n3179-88; as Nerine cirratulus by Hartman). San Francisco Bay, November 1951, 1 specimen (AHF n10172; as Nerinides acuta by Hartman). Hunter's Point, May 1972, 4 specimens (CAS 000701- 02). Oakland Inner Harbor, 1973-1974, 7 specimens (CAS 000705, 000707- 09, 000711-13). San Pablo Bay, February 1973, 1 specimen (CAS 000703). South San Francisco Bay, San Bruno Shoal, February 1973, 3 specimens (CAS 000704). Remarks.—This species was previously recorded in San Francisco Bay as Nerine cirratulus and Nerinides acuta by Hartman (1954) and as the latter species by Filice (1958) and Jones (1961). Foster (1971:62) commented on the great variability regarding both the configuration of the hooded hooks and their presence or absence in the posterior notopodia. The specimens from San Francisco Bay bear both bi- and tridentate hooded hooks, as well as worn unidentate ones (Figs. 2a-f, 3b), thus supporting Foster's observa- tions. In addition, the shape of the neuropodial lamellae in the middle and posterior regions is quite variable; they may or may not exhibit the notched ~l Ol VOLUME 90, NUMBER 1 and bilobed appearance which is presumed to distinguish the subgenus Scolelepis from the subgenus Nerinides (see Pettibone, 1963:90; Foster, 1971:59). This feature shows a continuous distribution within a single sample. On the basis of adult morphology I am unable to distinguish be- tween the paratype of Spio acuta from San Diego, the holotype of Nerine minuta from Texas, specimens identified as Nerine agilis from the north- east coast of North America (AMNH 1822), and examples of Scolelepis squamata from both coasts of North America, the Netherlands and Italy. This agrees with the observations of Foster (1971:59). Scolelepis squamata may be distinguished from S. tridentata (Southern ) and S. alaskensis (Treadwell) [see below] by having hooded hooks which are mainly bi- and tridentate when unworn (Fig. 2a—d), and which, even when worn, still show traces of the secondary tooth (Fig. 2e, f); they are first present in the neuropodia from setiger 25-40. The capillaries bear a well developed sheath and are strongly limbate with no trace of a distal fringe (Fig. 3a); the margin of the limbus is entire and has no ribbing. The primary hoods are more than 5-6 times the length of the main fang ( Fig. 3b). Scolelepis tridentata (Southern) Fig. 3c-e Nerinides tridentata Southern, 1914:98, pl. 10: fig. 23A-J.—Blake, 1975:214. Scolelepis (Nerinides) tridentata: Pettibone, 1963:93. Material—tIreland: Blacksod Bay, Clare Island Survey sta. W135, W160, W166, March & September 1910, from Laminaria roots, 3 syntypes (NMI 49.1910, 448.1910, 32.5.1914). California: Tomales Bay, Marin Co., 16 March 1957, 1 specimen (PMSM 180238; as Nerinides tridentata by Hartman). Pillar Point, San Mateo Co.., several specimens (examined courtesy of Moss Landing Marine Labora- tories ). Remarks.—The syntypes of Nerinides tridentata differ from the original description in several important respects. The hooded hooks are multi- dentate (Fig. 3d, e), with from 3-5 (usually 4 or 5) apical teeth surmounting the distinctly elongate main fang; they were erroneously described as tri- dentate by Southern (1914:99, Pl. 10, Fig. 23-J). Notosetae are present on setiger 1, contrary to Southern’s description (p. 98). Specimens from central California are identical in all respects to Southern’s types. Numerous larvae have been collected from Tomales Bay, Marin Co., California and success- fully raised to metamorphosis in the laboratory (J. A. Blake, personal com- munication ). Scolelepis tridentata is readily distinguished from S. squamata by bearing a short, stubby occipital cirrus between the bases of the palpi, by multi- dentate hooded hooks which first appear in the neuropodia from setiger 76 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON LENGTH OF MAIN FANG LENGTH OF HOOD 05 b Ay SS SS SSI Sa SS EEN SS HEN SS BANS SS ZN SS WN. LENGTH OF SS aw MAIN FANG = aw S= GNN LENGTH OF SS AN HOOD == RS 006 Naueacna SS FRONTAL Sea VIEW RS SSS I Sa 024 e ~ LATERAL | VIEW | a—b, Scolelepis squamata (Miller): a, Limbate capillary seta; b, Bidentate | neuropodial hooded hook in right lateral view, showing hood:main-fang ratio. c—e, Scolelepis tridentata (Souther), from syntypes of Nerinides tridenata: c, Limbate capillary seta, showing serrate margin and close-set ribbing; d, Multidentate neuropodial hooded hook in right lateral view, showing hood:main-fang ratio; e, Neuropodial hook with 5 apical teeth in latero-frontal aspect, hood removed. Scales in mm. oie, 8), 15-16, and by these hooks bearing hoods less than 2 times as long as the main fang (Fig. 3d). In addition, the capillary setae of S. tridentata, especially in the anterior setigers, exhibit a distinctly crenulate limbus bearing fine, closely set ribs (Fig. 3c). Scolelepis alaskensis (Treadwell) Figs. 4a-e, 5c Scolecolepis alaskensis Treadwell, 1914:201, pl. 12: figs. 21, 22. Scolelepis alaskensis: Pettibone, 1963:92. VOLUME 90, NUMBER 1 77 Fig. 4. Scolelepis alaskensis (Treadwell), from paratype of Scolecolepis alaskensis. a—d, Hooded hooks; a, Unworn hook; b, Worn hook, hood removed; c, Partly worn hook, hood laterally displaced; d, Worn hook; e, Alimbate capillary seta, showing distal fringe. Scales in mm. Material—Alaska, Shumagin, paratype (AMNH 770). Remarks.—In S. alaskensis, the hooded hooks begin in the neuropodia from setiger 57, and all are unidentate; when unworn, they exhibit a faleate, tapering point (Fig. 4a, c), or, when worn, bear a rounded or irregular knob- like tip with a terminal cavity (Fig. 4b, d). The capillary setae are alimbate and distally fringed (Fig. 4e). The pre- and postsetal lamellae in both rami of the middle and posterior regions form thick, glandular tori ( Fig. dc), quite unlike the protruding and non-glandular parapodia of S. squamata (see Foster, 1971, Figs. 122-124) or S. tridentata (see Southern, 1914, Fig. 23D-G ). Pettibone (1963:92) suggested that S. alaskensis may be reterrable to Scolelepis foliosa (Audouin & Milne-Edwards, 1833). Three specimens from Blacksod Bay, Ireland (NMI 283; Clare Island Survey sta. W115, Sep- 78 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON tember 1909) identified as Nerine [= Scolelepis] foliosa by Southern (1914: 100) were examined and found to be similar in all respects to S. alaskensis. However, a faint “gelatinous” sheath seems to be present around the distal timbriations of the capillaries in Southern’s material; it is not clear whether this might have been present originally, but subsequently worn away, in the paratype of S. alaskensis. The 2 species are otherwise indistinguishable. Dispio uncinata Hartman Spio setosa: Behre, 1950:13. [Not Verrill, 1873.] Dispio uncinata Hartman, 1951:87; 1961:88; 1969:105.—Foster, 1971:73.— Fauchald, 1973:24——Hodgson & Nybakken, 1973:220. Spio sp.: Hartman, 1954:10.—?Jones, 1961:316. Material—Calitornia: San Francisco Bay, west Berkeley beaches, July 1891, 1 specimen (AHF n3571; as Spio sp. by Hartman). Tomales Bay, Marin Co., June 1941, 5 specimens (AHF n294). Remarks.—New record for San Francisco Bay and Tomales Bay. This species was previously reported from San Francisco Bay as Spio sp. by Hart- man (1954) and possibly by Jones (1961). The material examined con- stitutes the first records for Dispio uncinata in this region, although the col- lection dates of 1891 and 1941 predate the original species description in 1951. It was recently reported from Monterey Bay by Hodgson & Nybakken (1973). J. A. Blake (personal communication) has recovered a number of larvae from plankton hauls in Tomales Bay, which upon metamorphosis proved to be D. uncinata. The species is otherwise known from southern California (Hartman, 1961), Massachusetts to North Carolina and the Carib- bean Sea (Foster, 1971), the Gulf of Mexico [type-locality], both coasts of Central America (Foster, 1971; Fauchald, 1973) and Chile (Foster, 1971). Foster (1971:76-78) has commented on the extreme variability in the num- ber of serrations on the anterior notopodial lamellae and on the number of setigers bearing such serrated lamellae, even between individuals from the same locality. Since these 2 features are the primary criteria separating the various species described for this genus, Foster (1971:77-78) noted that D. schusterae Friedrich, 1956 and D. remanei Firedrich, 1956, both from Central America, should probably be referred to D. uncinata. D. maroroi Gibbs, 1971, from the Solomon Islands, also falls within the range of vari- ability exhibited by D. uncinata with respect to these characters, and the status of the former species is unclear. Spiophanes berkeleyorum Pettibone Fig. 5a, b Spiophanes cirrata: Berkeley, 1927:12.—Hartman, 1954:10. [Not Sars, 1872. | VOLUME 90, NUMBER 1 79 Spiophanes sp.: Hartman, 1955:45. Spiophanes fimbriata: Hartman, 1966:214, 408 (part). [Not Moore, 1923.] Spiophanes berkeleyorum Pettibone, 1962:78.—Uschakov, 1971:1404. Material.—British Columbia: Departure Bay, east coast Vancouver Island, April 1936, 6 paratypes of S. berkeleyorum (USNM 30400). California: Off Redondo Beach Pier, 73 m, Velero IV sta. 2193-52, many specimens (AHF; as Spiophanes sp. by Hartman, 1955). Southern Cali- fornia, 7-15 m, Velero IV sta. 6105-59, 1 specimen plus fragments; Los Angeles—Long Beach outer harbor, Velero IV sta. 6107-59, 2 specimens, as Spiophanes fimbriata by Hartman (1966:214) [AHF]. San Francisco Bay, Oakland Inner Harbor, 9-12 m, 5 June 1974, 1 specimen (CAS 000013). Hunter's Point & Oakland Inner Harbor, 1974-1975, many specimens (CAS 001288-90, 001913). Monterey Bay, Elkhorn Slough, December 1975, 1 specimen (CAS 000706). Remarks.—The range of the species is here extended from Puget Sound to San Francisco Bay, Monterey Bay and Los Angeles Harbor. It was pre- viously reported from San Francisco Bay as Spiophanes cirrata by Hart- man (1954). Spiophanes berkeleyorum differs from the closely related S. kroeyeri [see below] in having a prominently bell-shaped prostomium with well developed lateral swellings at the anterior end, and in bearing highly developed, glandular, platelike lateral cephalic lobes with raised rims (Fig. 5a, b). There is no trace of the interparapodial genital pouches seen in S. kroeyeri (see also Pettibone, 1962:82, 86). Pettibone (1962:86) partly distinguished S. berkeleyorum and S. kroeyeri on the basis of differing num- bers of neuropodial hooks per ramus; however, this feature shows consider- able overlap between the two species and is not a useful diagnostic criterion. Spiophanes berkeleyorum is known in fairly shallow bays and harbors from the Strait of Georgia (British Columbia) and Puget Sound to southern California, and it is recorded from the Sea of Japan (Uschakoy, 1971). Both S. berkeleyorum and S. kroeyeri have been reported as Spiophanes fimbriata [see below], and it is not clear which records should be referred to which species. Spiophanes kroeyeri Grube Fig. 5d Spiophanes kroyeri Grube, 1860:88, pl. 5: fig. 1—Pettibone, 1962:S5 [synonymy ]. Spiophanes fimbriata Moore, 1923:179.—Hartman, 1960:117; 1969: 183, figs. ]—-4.—Fauchald, 1972b:199.—Blake, 1975:215. Spiophanes kroeyeri: Fauchald, 1972a:29. Material—Norway: Sognefjorden, 61°08’15’N, 05°45’30°E, 1272 m, May 1966, 9 specimens (AHF; identified by kK. Fauchald). 80 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON California: Monterey Bay, 697 m, Albatross sta. 4541, June 1904, holotype of S. fimbriata (USNM 17383). Off Los Angeles breakwater, 366 m, Velero IV sta. 2354-53, 1 specimen (AHF; as S. fimbriata by Hartman). From deep water off Mexico, sta. 18743-70, 18°12’03’N, 104°01’00’W, 2334 m, 1 speci- men (AHF F4095; as S. fimbriata by Fauchald, 1972b). Remarks.—Pettibone (1962:85) placed Spiophanes fimbriata Moore from 69-1158 m off central and southern California into synonymy with S. kroeyeri of cosmopolitan distribution [type locality, Greenland]. A com- parison of the holotype of S. fimbriata with specimens of S. kroeyeri from Norway supports this view. Spiophanes kroeyeri is readily distinguished from S. berkeleyorum [see above] by bearing prominent interparapodial genital pouches from setiger 14-15, in having a subtriangular prostomium without obvious lateral swellings, and in exhibiting reduced lateral cephalic lobes without raised, glandular rims (Fig. 5d). Specimens from deep water off Los Angeles, California and western Mexico identified as S. fimbriata by Hartman (1960) and Fauchald (1972b) agree in all respects with S. kroeyeri. Spiophanes kroeyeri appears to be a cosmopolitan, cold-water species frequently found in deep water off west- ern North America, and many of the records of S. fimbriata should prob- ably be referred to this species [see also remarks under S. berkeleyorum]. Spiophanes bombyx (Claparede ) Fig. 5e-¢ Spio bombyx Claparede, 1870:485. Spiophanes bombyx: Hartman, 1969:181, figs. 1-5.—Foster, 1971:40, figs. 66-75 [synonymy ].—Blake, 1975:215. Material —California, San Francisco Bay, off Alcatraz Island, September 1973 & June 1974, 6 specimens (CAS 0001914-15). Remarks.—New record for San Francisco Bay. The present specimens bear neuropodial hooded hooks which are bidentate (Fig. 5e), tridendate (Fig. 5£), or quinquedentate (Fig. 5g) with 3 very small superior teeth in a row overlying the apical tooth. Most authors report the presence of only bidentate hooks. The presence of a third tooth was also noted by Mesnil (1897:92) and Foster (1971:43). Prionospio (Prionospio) steenstrupi Malmgren Prionospio steenstrupi Malmgren, 1867:202.—Foster, 1971:84, figs. 175-185 [synonymy ]. Prionospio malmgreni: Berkeley, 1927:414.—Berkeley & Berkeley, 1952:29, figs. 54, 55—Hartman, 1969:159, figs. 1-4. [Not Claparéde, 1869. ] Material.—British Columbia, Pipers Lagoon, May 1920, 10 specimens (USNM 40794; as P. malmgreni by E. & C. Berkeley ). —E VOLUME 90, NUMBER 1 8] LOBE LATERAL CEPHALIC LOBE Fig. 5. ab, Spiophanes berkeleyorum Pettibone: a, Anterior end, dorsal view, setae and palpi not shown; b, Anterior end, ventral view, setae not shown. c, Scolelepis dlaskensis (Treadwell), from paratype of Scolecolepis alaskensis, right median para- podium; d, Spiophanes fimbriata Moore [= S. kroeyeri Grube], from holotype of S. fimbriata, anterior end, dorsal view, setae and palpi not shown; c—g, Spiophanes bombyx (Claparéde), neuropodial hooded hooks: e, Bidentate hook; f, Tridentate hook; g, Quinquedentate hook. Scales in mm. California: Velero IV sta. 2314-53, 3585-55, several specimens [AHF]. San Francisco Bay, off Angel Island, 22 m, February 1973, 1 specimen (CAS 000738 ). Remarks.—New record for San Francisco Bay. P. steenstrupi has been separated from P. fallax S6derstr6m, 1920 [= P. malmgreni: of authors (in part) ] by the absence of a high dorsal fold on setiger 7 or by exhibiting only very low folds on a variable number of segments. P. fallax, in contrast, 82 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON is said to bear a well developed transverse dorsal fold on setiger 7 (Sdder- strom, 1920:236; Foster, 1971:86-87, 89). Some specimens from western Canada possess a high dorsal fold on setiger 7 and subsequent segments (see also Berkeley & Berkeley, 1952:29; as P. malmgreni), whereas others in the same sample bear low dorsal crests. Foster (1971:87) thus noted that P. steenstrupi and P. fallax might ultimately prove to be conspecific. The single specimen from San Francisco Bay and other examples from southern California all exhibit moderately to well developed dorsal folds on a number of segments from setiger 7, and this character is regarded as variable. In view of the foregoing discussion, 2 paratypes of Prionospio cristata Foster, 1971 from North Carolina (USNM 43002) were examined. They do not differ to any appreciable extent from P. steenstrupi, and may prove to be juvenile forms of the latter. Prionospio (Apoprionospio) pygmaea Hartman Prionospio pygmaeus Hartman, 1961:93; 1969:381.—Reish, 1968:84.—Hodg- son & Nybakken, 1973:221.—Blake, 1975:215. Prionospio malmgreni: Hartman, 1955:77 (part). [Not Claparede, 1869. | Apoprionospio pygmaea: Foster, 1969:381; 1971:94, figs. 213-225 [syn- onymy ]. Material—California: San Francisco Bay, Alcatraz Island, September 1973, 3 specimens (CAS 001911). Central San Francisco Bay, June 1974, 1 specimen (CAS 002336). Outside Golden Gate Bridge, 37°46’00’N, 122°51’30’W, 3 specimens (CAS 000731). Remarks.—New record for San Francisco Bay. This species was recently recorded from Monterey Bay by Hodgson & Nybakken (1973). It is other- wise known from southern California (Hartman, 1961, 1969), the Gulf of California (Reish, 1968) and the Gulf of Mexico (Foster, 1971). It is com- mon in Tomales Bay, Marin Co. (J. A. Blake, personal communication ). Prionospio (Minuspio) cirrifera Wiren Prionospio (?) cirrifera Wiren, 1883:409. Prionospio cirrifera: Jones, 1961:266, 288, 316. Minuspio cirrifera: Foster, 1971:108, figs. 262-275 [synonymy]. Material—California: San Francisco Bay, off Potrero, February 1972, 2 specimens (CAS 000057). Off Angel Island, February 1973, 2 specimens (CAS 000786-87 ). Alcatraz Island, Oakland Inner Harbor, Hunter’s Point, September & December 1973, 11 specimens (CAS 001286-87, 001366, 001386, 001912). Remarks.—This species was previously reported from San Francisco Bay off Point Richmond by Jones (1961). None of the San Francisco Bay VOLUME 90, NUMBER 1 83 material exhibits any trace of the interparapodial pouches described for some specimens by Foster (1971:110). Paraprionospio pinnata (Ehlers ) Prionospio pinnata Ehlers, 1901:163. Prionospio alata Moore, 1923:185. [See also Foster, 1969:389. | Prionospio plumosa Treadwell, 1931:4 [homonym]. [Not Sars, 1872.] [See also Foster, 1969:389. ] Prionospio treadwelli Hartman, 1951:84 (part; not species from Louisiana ). Prionospio ornata Berkeley & Berkeley, 1961:660. Fide Foster, 1969:389. Paraprionospio pinnata: Foster, 1969:389, figs. 12-21; 1971:102, figs. 237- 246 [synonymy ].—Fauchald, 1972b:188.—Blake, 1975:213-214. Material_—California: Monterey Bay, 102 m, Albatross sta. D4549, holo- type of Prionospio alata (USNM 17369). Central San Francisco Bay, Feb- ruary 1973, 1 specimen (CAS 001907). Outside Golden Gate Bridge, 37°30’21”"N, 122°24’21’”W, August 1973, 1 specimen (CAS 001908). Maryland: Cheasapeake Bay, Fish Hawk sta. 8881, October 1920, holo- type of Prionospio plumosa Treadwell (USNM 19598). North Carolina: One specimen in vial with red cloth, presumably an original specimen used by Hartman (1951) in designating P. treadwelli as a replacement name for P. plumosa (AHF n1564). Remarks.—New record for San Francisco Bay. P. pinnata closely re- sembles P. krusadensis Fauvel, 1929 [type-locality, Gulf of Manaar, India] as described from Japanese material by Okuda (1937:244-246) and Imajima & Hartman (1964:284-285). However, the latter species does not bear strongly developed lateral peristomial expansions, the 3 pairs of branchiae begin on setiger 2 (the fused peristomium and following abranchial segment bear small neurosetae) and the hooded hooks are first present in the neuro- podia from setiger 16-17 (rather than from setiger 9 as in P. pinnata). Foster (1969:389) referred P. alata to P. pinnata; this was challenged by Fauchald (1972b:188). According to Moore (1923:185-186), P. alata bears branchiae on setigers 2-4 (“somites III, IV and V”) and hooded hooks are first found in the neuropodia from setiger 19 (“somite XX”). However, the holotype shows no evidence of setae prior to the first branchial seg- ment, and the neuropodial hooded hooks begin on setiger 9. ‘The synonymy of P. alata with P. pinnata is herein supported. Prionospio treadwelli was designated by Hartman (1951:85) as a replace- ment name for P. plumosa Treadwell. She described this species as bearing 4 pairs of branchiae, although Treadwell’s original description (1931:4) indicated that only 3 pairs were present. The holotype of P. plumosa from Cheasapeake Bay and Hartman’s specimen from North Carolina both bear 84 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 3, rather than 4, pairs of branchiae, and there is no evidence of any scars which would indicate the presence of additional gills. P. ornata Berkeley & Berkeley, described from larval material, was re- ferred to P. pinnata by Foster (1969:389). The presence in both P. krusadensis and P. ornata of a setigerous somite preceding the first branchial segment indicates the close affinity between Paraprionospio and Prionospio (sensu lato), as noted by Sdderstrom (1920). The genus Aquilaspio Foster, 1971 is here referred to the genus Paraprionospio Caullery, 1914. As noted by Foster (1971:101), Paraprionospio shows close affinities with the mono- typic genus Streblospio Webster, and the latter should probably also be considered one of the quasi-generic categories within the Prionospio-like group of spionids. Streblospio benedicti Webster Streblospio benedicti Webster, 1879a:120.—Hartman, 1954:10.—Filice, 1954: 16; 1958:191.—Jones, 1961:243.—Foster, 1971:112-113 [synonymy]. Streblospio lutincola Hartman, 1936:45, figs. 1, 2. Material—California: San Francisco Bay, north Berkeley beach, October 1932, several specimens (AHF n3453). Oakland Airport, November 1936, several specimens (AHF 3454). Berkeley beaches, July 1954, several speci- mens (AHF 3450). San Pablo Channel, Point San Pablo, August 1951, F. P. Filice, coll., several specimens (AHF n10396). Oleum, February 1972, several specimens (CAS 001371). South San Francisco Bay, off Redwood City, October 1973, 1 specimen (CAS 000714). Carquinez Strait, Mare Is- land Strait, Oakland Inner Harbor, south San Francisco Bay, Hunter's Point, Redwood City Harbor, 1973-1974, various stations, many specimens (CAS 001291-93, 001295-1305, 001909 ). Remarks.—This species is well known from San Francisco Bay (Hartman, 1936, 1954; Filice, 1954, 1958; Jones, 1961). Literature Cited Audouin, J. V., and H. Milne-Edwards. 1833. Classification des annélides et descrip- tion de celles qui habitent les cétes de la France. Ann. Sci. Nat. Paris 29:388— 412. Banse, K. 1972. On some species of Phyllodocidae, Syllidae, Nephtyidae, Goniadidae, Apistobranchidae, and Spionidae (Polychaeta) from the northeast Pacific Ocean. Pacific Sci. 26(20) :191-222. , K. D. Hobson, and F. H. Nichols. 1968. Annotated list of polychaetes. In U. Lie, A quantitative study of benthic infauna in Puget Sound. Fiskerdirekt. Skr. Ser. Havundersok. 14(5):521-548. Bamard, J. L. 1958. Amphipod crustaceans as fouling organisms in Los Angeles—Long Beach Harbors, with reference to the influence of seawater turbidity. Calif. Fish Game 44:161-170. VOLUME 90, NUMBER 1 85 Behre, E. H. 1950. Annotated list of the fauna of the Grand Isle region. Occ. Pap. Mar. Lab. Louisiana State Univ. 6:66 pp. Berkeley, E. 1927. Polychaetous annelids from the Nanaimo district. 3. Leodicidae to Spionidae. Contrib. Canad. Biol., N. S. 3(17) :407—422. , and C. Berkeley. 1936. Notes on Polychaeta from the coast of western Canada. 1. Spionidae. Ann. Mag. Nat. Hist., Ser. 10, 18:468—476. , ain 1952. Annelida. Polychaeta Sedentaria. In Canadian Pacific Fauna. Fish. Res. Bd. Canada 9b(2). Univ. Toronto Press, 139 pp. , and 1954. Additions to the polychaete fauna of Canada, with com- ments on some older records. Jour. Fish. Res. Bd. Canada 11(4) :454—471. , and 1961. Notes on Polychaeta from California to Peru. Canad. Jour. Zool. 39:659-664. Blake, J. A. 1966. On Boccardia hamata (Webster), new combination (Polychaeta, Spionidae ). Bull. So. Calif. Acad. Sci. 65:176—184. 1971. Revision of the genus Polydora from the east coast of North America (Polychaeta: Spionidae). Smithsonian Contrib. Zool. 75:32 pp. —. 1975. Phylum Annelida; Class Polychaeta. In R. I. Smith and J. T. Carlton, (eds), Light’s Manual: Intertidal Invertebrates of the Central California Coast. University of California Press, Berkeley: 151-243. Blake, J. A.. and K. H. Woodwick. 1971. A review of the genus Boccardia Carazzi (Polychaeta: Spionidae) with descriptions of two new species. Bull. So. Calif. Acad. Sci. 70:31—42. , and 1975. Reproduction and larval development of Pseudopolydora paucibranchiata (Okuda) and Pseudopolydora kempi (Southern) (Polychaeta: Spionidae). Biol. Bull. 149(1):109-127. Caullery, M. 1914. Sur les polychétes du genre Prionospio Mgn. Bull. Soc. Zool. Paris 39:355-361. Chapman, J. W., and J. A. Dorman. 1975. Diagnosis, systematics, and notes on Grandidierella japonica (Amphipoda: Gammaridea) and its introduction to the Pacific coast of the United States. Bull. So. Calif. Acad. Sci. 74:104—108. Claparéde, E. 1863. Beobachtungen tiber Anatomie und Entwicklungsgeschichte wirbel- loser Thiere an der Kiiste von Normandie angestellt. Leipzig, vii + 120 pp. —. 1869. Les annélides chétopodes du Golfe de Naples. Mém. Soc. Phys. Hist. Nat. Genéve 20(1):1-225. 1870. Les annélides chétopodes du Golfe de Naples. Mém. Soc. Phys. Hist. Nat. Genéve 20(2):365-542. Dean, D., and P. A. Hatfield. 1963. Pelagic larvae of Nerinides agilis (Verrill). Biol. Bull. 124(2):163-169. 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Invertebrates of the estuarine portion of San Francisco Bay. Wasmann Jour. Biol. 16:159-211. Foster, N. M. 1969. New species of spionids (Polychaeta) from the Gulf of Mexico and Caribbean Sea, with a partial revision of the genus Prionospio. Proc. Biol. Soc. Wash. 82( 28) :381—400. ——. 1971. Spionidae (Polychaeta) of the Gulf of Mexico and the Caribbean Sea. Stud. Fauna Curacao Caribb. Islands 36(129):183 pp. Freidrich, H. 1956. Mitteilungen tiber neue und wenig bekannte Polychaeten aus Mittel-und Siid-Amerika. Senckenbergiana Biol. 37:57-68. Gibbs, P. E. 1971. The polychaete fauna of the Solomon Islands. Bull. Brit. Mus. (Nat. Hist.); Zool. 21(5):99=211. Graham, H. W., and H. Gay. 1945. Season of attachment and growth of sedentary marine organisms at Oakland, California. Ecology 26:375-—386. Grube, A. E. 1860. Beschreibung neuer oder wenig bekannter Anneliden. Arch. Naturg. 26:71-118. Hartman, O. 1936. New species of Spionidae (Annelida: Polychaeta) from the coast of California. 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Part I. Preliminary results. Allan Hancock Pacific Exped. 19:1—185. 1956. Polychaetous annelids erected by Treadwell, 1891 to 1948, together with a brief chronology. Bull. Amer. Mus. Nat. Hist. 109(12):239-310. 1960. Systematic account of some marine invertebrate animals from the deep basins off southern California. Part II. Allan Hancock Pacific Exped. 22(2): 69-215. 1961. Polychaetous annelids from California. Allan Hancock Pacific Exped. 2322 0m pp: —. 1966. Quantitative survey of the benthos of San Pedro Basin, southern Cali- fornia. Part II. Final results and conclusions. Allan Hancock Pacific Exped. 19:187—456. 1969. Atlas of the Sedentariate Polychaetous Annelids from Califormia. Allan Hancock Foundation, University of Souther California, Los Angeles, 812 pp. Hobson, K. D. 1974. Orbiniella nuda new species (Orbiniidae) and nine new records of other sedentariate polychaetous annelids from Washington and British Colum- bia. Canad. Jour. Zool. 52:69-75. VOLUME 90, NUMBER 1 87 Hodgson, A. T., and J. W. Nybakken. 1973. A quantitative survey of the benthic in- fauna of northern Monterey Bay, California. Moss Land. Mar. Lab. Tech. Publ. 73-8:241 pp. Horst, R. 1920. Polychaete anneliden uit het Alkmaarder Meer door Dr. R. Horst. Zool. Meded. Leiden 5:110-111. Imajima, M., and O. Hartman. 1964. The polychaetous annelids of Japan. Part II. Allan Hancock Found. Publ. Occ. Pap. 23:239—452. Jacobi, R. 1883. Anatomisch-histologische Untersuchung der Polydoren der Kieler Bucht. Inaug. Dissert. Kiel, 35 pp. Jones, M. L. 1961. A quantitative evaluation of the benthic fauna off Point Richmond, California. Univ. Calif. Publ. Zool. 67(3) :219-320. Light, S. F. 1941. Laboratory and Field Text in Invertebrate Zoology. Stanford Uni- versity Press, viii + 232 pp. Light, W. J. 1969. Polydora narica, new species, and Pseudopolydora kempi cali- fornica, new subspecies, two new spionids (Annelida: Polychaeta) from central California. Proc. Calif. Acad. Sci. 36( 18) :531—550. ———. 1976. The San Francisco Bay Project identification manual series: a new approach to invertebrate keys. Syst. Zool. 25:236—242. Malmgren, A. J. 1867. Annulata Polychaeta Spetsbergiae, Gronlandiae, Islandiae et Scandinaviae hactenus cognita. Ofv. K. Vetensk. Akad. Forh. 24:127—235. Mesnil, F. 1897. Etudes de morphologie externe chez les annélides. II. Remarques complémentaires sur les Spionidiens. La familie nouvelle des Disomidiens. La place des Aonides (sensu Tauber, Levinsen). Bull. Sci. France Belg. 30:83—100. Moore, J. P. 1923. The polychaetous annelids dredged by the U.S.S. “Albatross” off the coast of southern California in 1904. IV. Spionidae to Sabellariidae. Proc. Acad. Nat. Sci. Philadelphia 75:179-259. Muller, O. F. 1806. Zoologica Danica sev Animalium Daniae et Norvegiae rariorum ac minus notorum, Descriptiones et Historia. Havniae 4:1—46. Okuda, S. 1937. Spioniform polychaetes from Japan. Jour. Fac. Sci. Hokkaido Imp. init, BG) seria Pettibone, M. H. 1954. Marine polychaete worms from Pt. Barrow, Alaska, with addi- tional records from the North Atlantic and North Pacific. Proc. U.S. Nat. Mus. 103 (3324) :203-356. ——. 1962. New species of polychaete worms (Spionidae: Spiophanes) from the east and west coast of North America. Proc. Biol. Soc. Wash. 75:77-88. ——. 1963. Revision of some genera of polychaete worms of the family Spionidae, including the description of a new species of Scolelepis. Proc. Biol. Soc. Wash. 76:89-104. Read, G. B. 1975. Systematics and biology of polydorid species (Polychaeta: Spioni- dae) from Wellington Harbour. Jour. Roy. Soc. New Zealand 5(4):395-419. Reish, D. J. 1954. Polychaetous annelids as associates and predators of the crustacean wood borer, Limnoria. Wasmann Jour. Biol. 12:223-226. —. 1955. The relation of polychaetous annelids to harbor pollution. Public Health Rep. 70:1168—1174. 1959a. An ecological study of pollution in Los Angeles—Long Beach Har- bors, California. Allan Hancock Found. Occ. Pap. 22:119 pp. 1959b. Ecology of Amphipoda and Polychaeta of Newport Bay, California. Allan Hancock Found. Publ. Occ. Pap. 21:106 pp. ——. 196la. The use of the sediment bottle collector for monitoring polluted marine waters. Calif. Fish Game 47:261-272. 1961b. A study of benthic fauna in a recently constructed boat harbor in southern California. Ecol. 42:84—91. 88 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 1963. Further studies on the benthic fauna in a recently constructed boat harbor in southern California. Bull. So. Calif. Acad. Sci. 62:23-32. 1964. A quantitative study of the benthic polychaetous annelids of Catalina Harbor, Santa Catalina Island, California. Bull. So. Calif. Acad. Sci. 63:86—92. ——.. 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. Reish, D. J., and H. A. Winter. 1954. The ecology of Alamitos Bay, California, with special reference to pollution. Calif. Fish Game 40:105-121. Sars, M. 1872. Diagnoser af nye annelider fra Christianiafjorden, efter Professor M. Sars’s efterladte Manuskripter. Forh. Vidensk. Selsk. Christiania 1871:406—417. [GO ke Sarssredal: Schmarda, L. K. 1861. Neue wirbelloser Thiere beobachtet und gesammelt auf einer Reise um die Erde 1853 bis 1857. I. Turbellarien, Rotatorien und Anneliden. Pt. 2. Leipzig, 164 pp. Soderstrom, A. 1920. Studien tuber die Polychaetenfamilie Spionidae. Dissert., Alquist and Wicksells, Uppsala, 286 pp. Southern, R. 1914. Archiannelida and Polychaeta. Clare Island Survey. Proc. Roy. Irish Acad. 31(47):1-160. ———. 1921. Fauna of the Chilka Lake. Polychaeta of the Chilka Lake and also of fresh and brackish waters in other parts of India. Mem. Indian Mus. 5:563-659. Treadwell, A. L. 1914. Polychaetous annelids of the Pacific coast in the collection of the Zoological Museum of the University of California. Univ. Calif. Publ. Zool. 13:175-234. —. 1931. Three new species of polychaetous annelids from Chesapeake Bay. Proc. U.S. Nat. Mus. 79:1-5. —. 1939. New polychaetous annelids from New England, Texas and Puerto Rico. Amer. Mus. Novitates 1023:7 pp. Uschakoy, P. V. 1971. Amphipacific distribution of polychaetes. Jour. Fish. Res. Bd. Canada 28:1403-1406. Verrill, A. E. 1873. Report upon the invertebrate animals of Vinyard Sound and the adjacent waters, with an account of the physical characters of the region. U.S. Fish Comm. Rept. Pt. 1:295-778. Webster, H. E. 1879a. Annelida Chaetopoda of the Virginia coast. Trans. Albany Inst. New York 9:202-272. 1879b. The Annelida Chaetopoda of New Jersey. Ann. Rep. New York State Mus. Nat. Hist. 32:101—128. Wirén, A. 1883. Chaetopoder fran Sibiriska Ishafvet och Berings Haf insamlade under Vega-Expeditionen 1878-79. Vega-Exped. Vetenskapliga Iakttagelser 2:383-428. Note.—After the manuscript was in press I was able to examine 3 speci- mens of Boccardia proboscidea from San Francisco Bay (AHF n3460, n3459). The occurrence of this species is thus confirmed. Department of Invertebrate Zoology, California Academy of Sciences, San Francisco, California 94118. PROG, BIOL, SOC. WASH: 90(1), pp. 89-98 A NEW GENUS OF PACIFIC ETELINAE (PISCES: LUTJANIDAE) WITH REDESCRIPTION Ob TAI YEE-SPHCIES William D. Anderson, Jr., Harry T. Kami, and G. David Johnson Etelis filamentosus was described by Fourmanoir (1970) from a speci- men obtained from the stomach of an Alepisaurus ferox collected off New Caledonia. Our examination of the holotype and four considerably larger specimens collected off the Hawaiian Islands shows that Fourmanoir’s species is not an Etelis, but represents a related, undescribed genus of eteline lutjanid. Yoshino and Araga (1975) described Etelis nudimaxillaris from two specimens collected off Okinawa. Based on our examination of the paratype of Yoshino and Araga’s species, we conclude that Etelis nudi- maxillaris is a junior synonym of Fourmanoirs Etelis filamentosus. The purposes of this paper are to describe the new genus and to redescribe the single species filamentosus. We used standard methods in making counts and measurements. Counting gill rakers on largest specimens was difficult, because it was hard to reflect the opercle far enough to see clearly the elements at anterior ends of upper and lower limbs, and because of the presence of elements intercalated within the main series of rakers. Intercalated elements were excluded easily from series counted where rakers were well developed, but because of similarity of size were not distinguished readily from rudiments counted. As a result, a degree of judgement was introduced into counting gill rakers. Material examined is in the collections of The Academy of Natural Sciences of Philadelphia (ANSP); Bernice P. Bishop Museum, Honolulu (BPBM); Museum National d’Histoire Naturelle, Paris (MNHN); Depart- ment of Biology, University of the Ryukyus, Naha, Okinawa (URB); and U.S. National Museum of Natural History, Washington, D.C. (USNM). Randallichthys, new genus Diagnosis.—A genus of Etelinae differing from all other genera of Lutjan- idae by the following combination of characters: a well-developed series of longitudinal bony ridges on lateral surface of maxillary; premaxillaries not protractile, fixed by a frenum; anterior tip of lower jaw considerably anterior to that of upper jaw when mouth closed; maxillary without scales; vomer and palatines with teeth; opercular spines poorly developed in young, becoming vestigial or disappearing in larger specimens; gill openings extending well forward of front of eye, as in Aphareus; dorsal fin con- tinuous, indented just anterior to junction of spinous and soft portions, but 90 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON not as deeply incised as in Etelis; ultimate dorsal soft ray slightly shorter than penultimate ray in young, becoming slightly longer than penultimate ray in older individuals; ultimate anal soft ray equal to or slightly longer than penultimate ray. Pectoral fin short, 55.0 to 66.0 percent of head length. Description—Mouth slightly oblique, terminal. No fleshy protrusion at anterior end of upper lip. No teeth on pterygoids or tongue. Two nostrils on each side, close to each other and to eye. Interorbital region flattened, but not as much as in Aprion or Etelis—more like that of Aphareus. Gill arches four, a slit behind the fourth. Gill rakers armed with numerous conical teeth; lateral gill rakers on anterior gill arch with teeth confined almost exclusively to their medial surfaces. Pseudobranchiae present. Spines of dorsal and anal fins rather weak. Caudal fin lunate. Scales ctenoid. Cheek, postorbital region, opercle, subopercle, interopercle, and proximal concave border of preopercle with scales. A patch of scales in temporal region separated from postorbital scales below and dorsolateral scales of body above by narrow naked areas running obliquely posteriorly and ventrally from scaleless interorbital region; lower oblique naked area usually wider than upper; upper naked area usually crossed by bridge of scales at about mid-length, lower naked area bridged at its posterior end by scales; temporal patch of scales usually separated posteriorly from scales of opercle by naked area. Snout, preorbital, interorbital, a narrow zone below and behind eye, most of preopercle, maxillary, lower jaw, and gular region naked. Dorsal and anal fins scaleless. Pelvic axillary process present. Pec- toral and caudal fins scaly basally. Lateral line complete, sensory tubules simple. Pelvic-fin rays L5. Principal caudal-fin rays 17 (9 + 8); procurrent caudal rays 11 or 12 dorsally, 10 or 11 ventrally. Branchiostegal rays 7 (5 on ceratohyal + 2 on epihyal). Vertebrae 24 (10 precaudal + 14 caudal). Caudal with no hypural fusion. Epipleural ribs 8. Trisegmental pterygio- phores 5 to 6 dorsally and 4 to 5 ventrally. Three predorsal bones, three anteriormost dorsal pterygiophores, and four anteriormost neural spines in typical eteline configuration (first predorsal bone anterior to first neural spine, second and third predorsal bones between first and second neural spines, first dorsal pterygiophore between second and third neural spines, and second and third dorsal pterygiophores between third and fourth neural spines). Other characters are those in the generic diagnosis and those of the single species. Etymology.—The name Randallichthys is for John E. Randall of the Bernice P. Bishop Museum, Honolulu, Hawaii, in recognition of his numerous contributions to the understanding and appreciation of the biology of Indo- Pacific fishes; the suffix—ichthys (= fish)—is from the Greek. The gender is masculine. Type species.—Etelis filamentosus Fourmanoir, 1970. VOLUME 90, NUMBER 1 91 Randallichthys filamentosus (Fourmanoir) Migselee2e) alles ie 2 Etelis carbunculus: Fowler (non Cuvier, 1828), 1928:193-194, pl. XVII, A (synonymy referring to species of Etelis; description, in part, of Randall- ichthys filamentosus and in part, apparently, of a species of Etelis; illus- tration of Randallichthys filamentosus; Hawaii). Etelis filamentosus Fourmanoir, 1970:26, 28-29, fig. 6 (original description and illustration; holotype MNHN 1970-33, 136 mm SL; type-locality 2220279, 162° 20'E ) . Etelis nudimaxillaris Yoshino and Araga, 1975:236, pl. 61-H (original de- scription and illustration; holotype URB 78-0110, 424 mm SL; type- locality off Okinawa). Description.—Variable meristic data and selected morphometric data are in Tables 1 and 2. Dorsal-fin rays X,11. Anal-fin rays III,9. Lateral surface of maxillary with a series of ca. 4-9 longitudinal bony ridges (ca. 4 on holotype, ca. 6-9 on larger specimens); ridges, for the most part, present only on exposed portion of maxillary. Maxillary ending posteriorly some- what short of vertical through middle of eye. Middle of eye approximately on horizontal line through anterior end of snout. Posterior border of anterior nostril with flap of tissue reaching, when reflected, to or slightly past anterior border of posterior nostril. Anterior nostril elliptical, with dorso-ventral axis longer; posterior nostril horizontally elliptical to almost circular, larger than anterior nostril. Posteriormost point of head ending well anterior of vertical through base of first dorsal spine. Premaxillary of holotype with inner band of small, essentially villiform teeth, some of teeth at anterior end of band enlarged and recurved; outside of this band a series of conical teeth (progressively larger anteriorly), one tooth on each side near front of jaw enlarged as a small canine; symphysis toothless. Premaxillary den- tition of other specimens similar to that of holotype except no canines on three largest specimens. Dentary of holotype with series of conical teeth on rear and middle portions, anteriorly this series expanded into a band of essentially villiform teeth, some of which enlarged and recurved; outside of this band on each side two small exserted canines, one near symphysis and One on side of jaw at some distance posterior to symphysis; symphysis toothless. Dentary teeth of other specimens in similar pattern but all very small (villiform anteriorly) except for 3-7 exserted conical teeth on each side, in series starting near symphysis and extending posteriorly on side of jaw. Larger teeth on premaxillaries and dentaries needle sharp. Vomer and palatine with small conical teeth on holotype (villiform on other specimens); those on vomer in roughly chevron-shaped patch with apex blunt and directed anteriorly; teeth on palatine in narrow antero-posterior|) oriented band. Preopercle serrate but without spine at angle; holotype with 92 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Table 1. Meristic data for Randallichthys filamentosus. Holotype is MNHN 1970-33; paratype of Etelis nudimaxillaris is URB 78-0111. Standard length is in mm. For bilateral counts the left side is presented first. MNHN BPBM URB USNM BPBM ANSP Character 1970-33 19695 78-0111 216255 11365 95004 Standard length 136 242, A408 A87 A493 499 Pectoral-fin rays 16,16 IAG 16,16 16,16 Sly 16,16 Gillrakers (including rudiments ): Upper limb 9.8 7,8 6,5 8,8 el 8,7 Lower limb 14,14 1S ISS 14,14 14,14 14,16 14,14 Total Ore, TOE e 20,19* ZOE? eS DEON! Tubed lateral-line scales 49,49 48,49 49, 49 49 49 49 48 49 49 Scales above lateral line —— (9) 9,10 ca.8,9 8, ca.9 — 8,9 Scales below lateral line —= .118} Le 14,13 13,14 4, — 14,14 Cheek-scale rows =e ONCanOe, sCane.9 Sis) = OL, SS 9,9 Predorsal scales ca. 20 24 ca. 23 ca. 22 ca. 22 cael Caudal-peduncle scales 2, 24 22 23 2 23 Scale rows between lateral line & middorsal fin 5.0,— 6:5,6.5,°'6.0:6.0'- 65:65) GiOiGIOR GG * The anteriormost gill arch was removed on each side. Although it appears that all gill rakers were removed intact along with the arches, there is a possibility that some were not. both limbs serrate, serrae larger in region of angle; other specimens with small serrae on horizontal limb, at angle, and on vertical limb near angle, preopercular notch and margin of vertical limb above notch smooth to almost smooth. Preopercular notch slightly developed only on one side of holotype, slightly to well developed on other specimens. Holotype with margins of interopercle and subopercle serrate for some distance on either side of their junction; other specimens without serrae on margins of inter- opercle and subopercle. Body compressed, not particularly deep, shape resembling that of Aphareus or Etelis. Pectoral fin short, reaching a vertical from near base of seventh dorsal spine on holotype and a vertical through base of eighth or ninth dorsal spine on other specimens. Two dorsal- most and one or two ventralmost pectoral-fin rays unbranched, other pectoral-fin rays branched. Pelvic fin ending anterior to vent, reaching to a vertical from near base of ninth dorsal spine. Lower lobe of caudal fin with some rays excessively produced and filamentous on holotype (not produced on other specimens). Leading edge of anal fin forming almost a right angle with distal edge, anal fin frequently slightly angulated posteriorly. Predorsal scales beginning over posterior part of orbit. Rows of scales above and below lateral line parallel to it. Largest specimens quite oily. VOLUME 90, NUMBER 1 93 Table 2. Morphometric data for Randallichthys filamentosus. Holotype is MNHN 1970-33; paratype of Etelis nudimaxillaris is URB 78-0111. Standard length in mm, other measurements in percentage of standard length. MNHN BPBM URB USNM BPBM ANSP Character ISOS Sree LOO Io mn O- Olas G2 ys 11365 95004 Standard length 136 242, 408 487 493 499 Head length 30.8 ole 31.0 30.2 30.9 30.1 Snout length 9.9 9.6 9.8 NOR 10.0 10.0 Fleshy orbit, horizontal diameter 7.6 8.4 C5 6.8 6.8 6.6 Interorbital, least bony width Toll UP eo 8.2 7.8 7.9 Suborbital, least width 4.1 4.0 3.9 3.9 4.0 3.6 Upper jaw length IBA IBsT 14.0 13.8 Sel 13.4 Lower jaw length 16.8 en ie 16.5 16.0 16.8 Body depth at first dorsal spine 31.2 30.7 31.0 32.0 Bondl 32.1 Body greatest width Ge 13.0 15.9 16.8 16.5 16.0 Caudal peduncle, least depth 11.7 ON ILL AL IES 10.5 11.4 Penultimate dorsal soft ray length ca. 10.4 8.6 8.6 Tess Bn TD eo Ultimate dorsal soft ray length CanlOnl 8.4 8.9 7.4 er 7.6 Penultimate anal soft ray length 9.4 COR CaM ES Toll 6.5 eal Ultimate anal soft ray length 9.4 Tol 8.6 Dole 6.7 el Pectoral fin length 16.9 19.3 18.8 WS 18.8 19.8 Pelvic fin length PL 21.0 5) 20.5 20a 20.6 Upper caudal-fin lobe length 27.5+ ca. 30.7 — SID — one Lower caudal-fin lobe length 83.1-— ca. 32.2 = 32.8 == 32.9 Coloration.—Fourmanoir (1970) stated that in formalin the holotype (MNHN 1970-33, 136 mm SL) showed four or five dark bands. These vertical bands (or bars) of dark pigmentation, although quite faded and not distinct in Fig: 1, are still evident on the specimen more than 15 years after preservation. The first of these on head immediately posterior to orbit, the second beneath spinous dorsal, the third beneath posterior spinous dorsal and anterior soft dorsal, and the fourth beneath soft dorsal; hint of a fifth bar on caudal peduncle. Bars more evident dorsally than ventrally. Base of dorsal fin, interradial membranes of spinous dorsal, proximal parts of interradial membranes of soft dorsal, interradial membranes of pelvic fins, and base of caudal fin with considerable dusky pigment. Anal fin lightly pigmented. Pectoral fins unpigmented. According to M. Legand, who collected the holotype, the coloration before preservation was ros) without dark bars (Fourmanoir, 1970). In alcohol a larger specimen (BPBM 19695, 242 mm SL; Fig. 2) is Y OF WASHINGTON aon i OCICALT SOE iOL R iv Py au OF TH INGS D 4 PROCEE 94 AGS) WU OFT “€-OL61 ‘(d1OUBUINO ) snsojyuawopnf shyyyoyjopupy Fo edA}o/OH ico VOLUME 90, NUMBER 1 95 mainly gray (darker dorsally) with darker bars placed similarly to those described for the holotype. Spinous dorsal and proximal part of soft dorsal fins heavily pigmented; narrow black band on spinous dorsal fin distally. Anterior two-thirds of anal fin lightly pigmented. Distal portion of soft dorsal fin, posterior one-third of anal fin, and pectoral fins unpigmented or essentially so. Right pelvic fin (left missing) heavily pigmented with broad black border distally. Upper and lower lobes of caudal fin with considerable pigment proximally, paler distally; middle of caudal fin more lightly pigmented proximally than upper and lower lobes, much paler distally. A color transparency of this specimen shows dark gray on dorsum of snout and on interorbital region; preorbital region, lips, and ventral part of lower jaw orange; postorbital region gray with pink to orange admixture; most of remainder of head pale pink; iris mainly orange. Body dull orange gray dorsolaterally, pale pink laterally and ventrally; dark vertical bars visible, but not very distinctive. Spinous dorsal fin bordered distally by narrow black band separated from general yellowish ground color of fin anteriorly by wider gray zone; soft dorsal fin yellowish. Anal fin yellow orange proximally with narrower pale zone distally. Pectoral fin pinkish. Pelvic fin yellow orange proximally with broad black margin distally. Caudal fin with dark gray blotch on each lobe near base, blotches connected by narrow gray line; most of caudal fin dull orange; most of distal edge of caudal fin dark. In alcohol, the four largest specimens lack any distinctive coloration on head or body, but interradial membranes of spinous dorsal fin with narrow dark distal edging. Anterior anal soft rays with some dark pigment distally. A dark band over distal margin of pelvic fin. Very fine dark distal edging on middle rays of caudal fin of two of largest specimens. In color prints of one of largest specimens (BPBM 11365, 493 mm SL) head and iris rosy. Ground color of body rosy with suffusion of yellowish pigment dorsolaterally. Dorsal fin grayish except: distal edges of inter- radial membranes of spinous dorsal fin black, rosy spot near distal margin of spinous dorsal fin about two-thirds distance from anterior end, and distal portion of posterior half of soft dorsal fin with yellowish stripe. Anal fin rosy with an admixture of yellow. Pectoral fin rosy proximally with some yellowish pigment distally. Leading edge of pelvic fin mostly rosy, most of fin yellowish, distal edge black. Caudal fin rosy with considerable yel- low influence in ground color, bright yellow band (resembling the numeral! seven) beginning proximally on upper rays (as horizontal arm of seven), proceeding distally to near posterior tip of fin and thence downward (as vertical arm of seven) parallel to posterior border, and ending some distance from ventral border of lower lobe; a narrow dark border along distal edge. A color print of another specimen (USNM 216255, 487 mm SL.) and the PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 96 ‘(posioAdl oATzesou “poydeisojyoyd opis }YSII) TS UW ZPZ “CBOBI Wadd ‘(Alourutino,, ) snsoquawppyf shysYyoujopupy ‘G ‘Bly VOLUME 90, NUMBER 1 97 illustration provided by Yoshino and Araga (1975, pl. 61-H) of Etelis nudimaxillaris resemble the color prints of the specimen described above. Synonymy.—Yoshino and Araga (1975) described Etelis nudimaxillaris from two specimens collected off Okinawa. Although there are several discrepancies between Yoshino and Araga’s description of Etelis nudi- maxillaris and our description of Randallichthys filamentosus, examination of the paratype of E. nudimaxillaris shows it to be conspecific with R. fila- mentous. Because the paratype of E. nudimavillaris agrees closely with the other specimens of R. filamentosus and because the illustration accompany- ing the original description of E. nudimazxillaris is a good likeness of R. filamentosus, we consider Etelis nudimaxillaris to be a junior synonym of Randallichthys filamentosus. Yoshino and Araga (1975) stated that Fowler (1928) figured and de- scribed their new species as Etelis carbunculus. Fowler's illustration (1928, pl. XVII, A) and his description, in part, are of R. filamentosus. Distribution.—This species is known from off the Hawaiian Islands, New Caledonia, and Okinawa in depths of 152-293 m. Material examined.—We examined 6 specimens, 136-499 mm SL. Holotype of Etelis filamentosus: MNHN 1970-33 (136 mm SL); off New Caledonia, west of Nouméa (well offshore), 22°52’S, 162°20’E; coll. M. Legand, 4 July 1961; from stomach of an Alepisaurus ferox caught by Japa- nese long line. Paratype of Etelis nudimaxillaris: URB 78-0111 (408 mm SL); Ryukyu Islands, off Okinawa; April 1973; hand line. Other material: BPBM 11365 (1 specimen, 493 mm SL); Hawaiian Islands, Hawaii (west coast), off Napoopoo; 293 m; coll. Weston Leslie, 25 June 1971; hook and line, 6pelu ( Decapterus pinnulatus ) as bait. USNM 216255 (1, 487 mm SL); Hawaiian Islands, Oahu, Tamashiro Fish Market: coll. J. E. Randall, 21 November 1975. ANSP 95004 (1, 499 mm SL); same data as USNM 216255 except 18 December 1975. BPBM 19695 (1, 242 mm SL); Hawaiian Islands, Oahu (west coast), off Waianae; 152 m; coll. John Rivera, 21 January 1976; hook and line. Distinguishing Characteristics and Relationships In addition to the characters given in the generic diagnosis, those useful for distinguishing Randallichthys filamentosus from other lutjanids are: dorsal-fin rays X,11; anal-fin rays III,9; anterior soft rays of dorsal and anal fins not elongated; dorsal and anal fins scaleless; lower lobe of caudal fin with some rays excessively produced and filamentous in young; no fleshy protrusion at anterior end of upper lip; anterior and posterior nostrils close to each other and to eye; interorbital region flattened; molaritorm teeth absent; pterygoids and tongue toothless. 98 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON The structure of the frontal region of the skull (i.e., presence of a posterior frontal thickening forming a complete transverse ridge of demarcation anterior to occipital region), the number of dorsal-fin rays, the configuration of the predorsal bones, and the possession of a well-developed accessory subpelvic keel all support the placement of Randallichthys in the lutjanid subfamily Etelinae. Randallichthys appears to be most closely related to Aphareus. It shares with that genus the following characters not found in combination in other eteline genera: premaxillaries not protractile, max- illary without scales, lower jaw extending well anterior to upper jaw, gill openings extending considerably anterior to orbit, and section A, of adductor mandibulae without an anterodorsal extension. Acknowledgments M. L. Bauchot, MNHN, J. E. Randall, BPBM, and T. Yoshino, URB, loaned us the specimens examined, and Randall allowed us to deposit specimens he obtained in the collections at ANSP and USNM. E. N. Gramblin, J. F. McKinney, and G. C. Van Dyke, USNM, and B. M. Martin, Medical University of South Carolina, made the radiographs used. Photo- graphs for Figs. 1 and 2 were provided by J. L. Russo, National Marine Fisheries Service Systematics Laboratory (NMFSSL), USNM. P. Kawamoto, Hawaii Fish and Game Division, and J. E. Randall made the photographs studied in writing descriptions of coloration of freshly collected specimens. Curators of the Division of Fishes, USNM, allowed us the use of space and facilities. Cindy Simmonds, Chesapeake Biological Laboratory (CBL), typed the manuscript, and D. M. Cohen, NMFSSL-USNM, W. L. Fink, USNM, and R. A. Fritzsche, CBL, reviewed it. This is contribution number 39 of the Grice Marine Biological Laboratory and contribution number 88 of the Marine Laboratory, University of Guam. Literature Cited Cuvier, G. 1828. In Cuvier, G., and A. Valenciennes. Histoire naturelle des poissons. 2. F. G. Levrault, Paris, xxi + 490 pp. Fourmanoir, P. 1970. Notes ichtyologiques (I). Cah. O.R.S.T.O.M., sér. Océanogr. 8(2):19-33, 10 figs. Fowler, H. W. 1928. The fishes of Oceania. Mem. Bernice P. Bishop Mus. 10, iii + 540 pp., 82 text figs., 49 pls. Yoshino, T., and C. Araga. 1975. In Masuda, H., C. Araga, and T. Yoshino. Coastal fishes of southern Japan. Tokai Univ. Press, Tokyo, 379 pp., 151 pls. Grice Marine Biological Laboratory, College of Charleston, 205 Fort Johnson, Charleston, South Carolina 29412; Marine Laboratory, University of Guam, Agana, Guam 96910; and Scripps Institution of Oceanography, University of California, San Diego, La Jolla, California 92037. PROC. BIOL. SOC. WASH. 90(1), pp. 99-107 A SYNOPTIC LIST OF THE DESCRIBED GROUND BEETLE LARVAE OF NORTH AMERICA (COLEOPTERA: CARABIDAE) Raymond G. Thompson Adequate descriptions and/or illustrations of 156 species and 3 subspecies of larvae of North American Carabidae have been published to date. This represents ca. 7% of the approximately 2,000 known ground beetle species occurring in North America. Reliable species identifications of these forms are difficult at best; taxonomic keys are few and the available descriptions widely dispersed in the literature. Kirk (1972) via a pictorial key made it possible to identify some common larvae found in South Dakota croplands. A few more species identifications are possible with the keys presented by Chu (1945), but some determinations accepted by Chu have not been associated with the adult forms and are, therefore, questionable. Other identification keys can be found in Burgess and Collins (1917) for the genus Calosoma (Carabini), Larsson (1941) primarily for larvae in Den- mark but including some holarctic and introduced species, and Lindroth (1954) for the genera Elaphrus and Blethisa (Elaphrini) which include some North American species. Kirk (1972) pointed out that accurate larvae-adult associations are possible by only 2 methods: first, by rearing field-collected larvae to the adult stage and then identifying the adults; second, by obtaining eggs and subsequent larvae from known adults brought into the laboratory. In recent years interest in studies of the immature stages of insects, in- cluding Coleoptera, has increased. The following list is presented in an effort to partially organize references concerning larval Carabidae, to bring attention to areas that need further work, and to stimulate interest in this group of beetles. This list includes only species of larvae which have verbal descriptions and/or usable illustrations. For purposes of this list, a verbal description is considered adequate if the larva is characterized sufficiently to distinguish it from similar larvae. An illustration is considered adequate if it is not too diagrammatic and can be used to distinguish the larva from similar larvae. Descriptions of larval species which have been synonymized and described subspecies are indicated. Larval species mentioned in the litera- ture as “known” without descriptions and/or illustrations are excluded. Finally, illustrations or verbal descriptions which have been presented with- out a species name are also excluded. 100 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Subfamily TRACHYPACHINAE Tribe TRACHYPACHINI Trachypachus gibbsi LeConte (Lindroth, 1960) Tribe GEHRINGINI Gehringia olympica Darlington (Lindroth, 1960) Subfamily CARABINAE Tribe CARABINI Carabus chamissonis Fisher (Lindroth, 1955a) Carabus limbatus Say (Burgess and Collins, 1917) Carabus maeander Fisher (Lapouge, 1929-32; Lindroth, 1954) Carabus nemoralis Mueller (Larsson, 1941) (syn) auratus Heuer (Schiodte, 1867, 1872) (syn) hortensis Fabricius (Larsson, 1941) Carabus serratus Say (Lindroth, 1954) Calosoma cancellatum Eschscholtz (Burgess and Collins, 1917; Larsson, 1941 ) Calosoma calidum Fabricius ( Burgess and Collins, 1917; Kirk, 1972) Calosoma externum Say (Burgess and Collins, 1917; Salle, 1849) Calosoma frigidum Kirby (Burgess and Collins, 1917) Calosoma haydeni Horn (Burgess and Collins, 1917) Calosoma marginalis Casey (syn) lugubre LeConte (Burgess and Collins, 1917) Calosoma obsoletum Say (Burgess and Collins, 1917) Calosoma peregrinator Guerin (Burgess and Collins, 1917) Calosoma scrutator Fabricius (Burgess and Collins, 1917) Calosoma semilaeve LeConte (Burgess and Collins, 1917) Calosoma sycophanta Linnaeus (Burgess and Collins, 1917; Larsson, 1941) Calosoma wilcoxi LeConte (Burgess and Collins, 1917) Tribe CYCHRINI Sphaeroderus lecontei Dejean (Lindroth, 1954) Sphaeroderus nitidicollis Chevrolat (Lindroth, 1954) Scaphinotus elevatus Fabricius (Wickham, 1893b) Tribe OMOPHRONINI Omophron americanum Dejean (Silvey, 1936) Omophron labiatum Fabricius (Schiodte, 1867, 1872) Omophron tesselatum Say (Silvey, 1936) VOLUME 90, NUMBER 1 101 Tribe NEBRIINI Pelophila borealis Paykull (Johnson and Carpenter, 1898; Andersen, 1970) Nebria brevicollis Fabricius (Schiodte, 1867, 1872; Larsson, 1941) Nebria diversa LeConte (syn) livida LeConte (Schiodte, 1867, 1872; Larsson, 1941 ) Nebria gyllenhali Schoenherr (Larsson and Gigja, 1959; Andersen, 1970) Nebria nivalis Paykull (Larsson and Gigja, 1959; Andersen, 1970) Tribe OPISTHIINI Opisthius richardsoni Kirby (Lindroth, 1960) Tribe NOTIOPHILINI Notiophilus aquaticus Linnaeus (Schiodte, 1867, 1872; Larsson, 1941) Notiophilus biguttatus Fabricius (Schiodte, 1867, 1872; Emden, 1942; Lars- son, 1941) Notiophilus novemstriatus LeConte (Thompson and Allen, 1974) Tribe LORICERINI Loricera pilicornis Fabricius (Schiodte, 1867, 1872; Bryson and Dillon, 1941) Tribe ELAPHRINI Diacheila arctica Gyllenhal (Lindroth, 1954) Blethisa julii LeConte (Lindroth, 1954) Blethisa multipunctata Linnaeus (Larsson, 1941) (sbsp) aurata Fischer (Lindroth, 1954) Blethisa quadricollis Haldman (Lindroth, 1941) Elaphrus clairvillei Kirby (Lindroth, 1954) Elaphrus lapponicus Gyllenhal (Lindroth, 1954 ) Elaphrus olivaceus LeConte (Lindroth, 1954) Elaphrus riparius Linnaeus (Schiodte, 1867, 1872; Larsson, 1941; Lindroth, 1954) Tribe BROSCINI Miscodera arctica Paykull (Andersen, 1968 ) Tribe SCARITINI Pasimachus elongatus LeConte (Kirk, 1972; Thompson and Allen, 1974) Scarites subterraneus Fabricius (Wickham, 1893b ) (sbsp) quadriceps Chaudoir (Emden, 1942) (syn) substriatus Haldeman (Kirk, 1972) 102 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Dyschirius interger LeConte (Kirk, 1972) Dyschirius politus Dejean (Larsson, 1941) Dyschirius sphaericollis Say (Silvey, 1936) Clivina fossor Linnaeus (Boving, 1910, 1911; Emden, 1942) Clivina impressifrons LeConte (Phillips, 1909; Kirk, 1972) Tribe BEMBIDIINI Tachyta nana Gyllenhal (Emden, 1942) Elaphropus anceps LeConte (Kirk, 1972) Elaphropus incurvus Say (Emden, 1942) Elaphropus parvulus Dejean (Cerruti, 1939) Elaphropus vivax LeConte (Emden, 1942) Bembidion confusum Hayward (Silvey, 1936) Bembidion constrictum LeConte (syn) contractum Dejean (Lindroth, 1955a) Bembidion guttula Fabricius (Larsson, 1941) Bembidion lampros Herbst (Larsson, 1941) Bembidion nitidum Kirby (Dimmock and Knab, 1904; Kirk, 1972) Bembidion quadrimaculatum Linnaeus (Kirk, 1972) Bembidion sulcipenne Sahlberg (syn) lenense Poppius (Lindroth, 1955a ) Bembidion tetracolum Say (Larsson, 1941; Kirk, 1972) Tribe POGONINI Thalassotrechus barbarae Horn (Moore, 1956) Tribe TRECHINI Trechus obtusus Erichson (Jeannel, 1920; Boldori, 1931; Boldori, 1932; Lars- son, 1941) Trechus rubens Fabricius (Larsson, 1941) Tribe PTEROSTICHINI Agonum albipes Fabricius (syn) ruficorne Goeze (Kemner, 1913) Agonum consimile Gyllenhal (Lindroth, 1955b ) Agonum extensicolle Say (Schaupp, 1881b; Dimmock and Knab, 1904) Agonum muelleri Herbst (Larsson, 1941) Agonum placidum Say (Kirk, 1972) Agonum punctiformis Say (Thompson and Allen, 1974) Agonum thoreyi Dejean (Larsson, 1941; Lindroth, 1955b ) Pterostichus adstrictus Eschscholtz (Goulet, 1974) VOLUME 90, NUMBER 1 103 Pterostichus adoxus Say (Dimmock and Knab, 1904) Pterostichus chalcites Say (Kirk, 1972) Pterostichus femoralis Kirby (Dogger and Olson, 1966) Pterostichus lucublandus Say (Schaupp, 188la; Wickham, 1893b) Pterostichus madidus Fabricius (Larsson, 1941; Emden, 1942 ) Pterostichus melanarius Illiger (Schiodte, 1867, 1872) Pterostichus mutus Say (Schaupp, 1881a) Pterostichus pensylvanicus LeConte (Goulet, 1974) Pterostichus strenuus Panzer (Larsson, 1941) Pterostichus stygicus Say (Dimmock and Knab, 1904) Pterostichus vernalis Panzer (Larsson, 1941) Abacidus permundus Say (Kirk, 1972) Abax parallelepipedus Piller & Mitterpacher (syn) ater villers (Kemner, 1913; Emden, 1942) (syn) striola Fabricius (Schiodte, 1867, 1872) Evarthrus alternans Casey (Kirk, 1972) Evarthrus torvus LeConte (Kirk, 1972) Synuchus impunctatus Say (Larsson and Gigja, 1959; Lindroth, 1956 ) Pristonychus terricola Herbst (Boving and Craighead, 1931) Calathus advena LeConte (Larsson and Gigja, 1959 ) Calathus fuscipes Goeze ( Boving, 1910, 1911; Kirk, 1972) Calathus melanocephalus Linnaeus (Kirk, 1972; Larsson and Gigja, 1959) Tribe AMARINI Amara aulica Panzer (Larsson, 1941) Amara brunnea Gyllenhal (Larsson, 1941) Amara carinata LeConte (Kirk, 1972) Amara cupreolata Putzeys (Dogger and Olson, 1966) Amara exarata Dejean (Kirk, 1972) Amara familiaris Duftschmidt (Larsson, 1941) Amara farcta LeConte (Kirk, 1972) Amara fulva Mueller (Larsson, 1941) Amara obesa Say (Lugger, 1899; Kirk, 1972) Tribe HARPALINI Trichocellus cognatus Gyllenhal (Larsson, 1941) Bradycellus rupestris Say (Chu, 1945) Stenolophus comma Fabricius (Kirk, 1972) Stenolophus conjunctus Say (Chu, 1945) Stenolophus lineola Fabricius (Chu, 1945) Stenolophus pallipes Fabricius (Bryson and Dillon, 1941) Acupalpus meridianus Linnaeus (Larsson, 1941) 104 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Notiobia picea LeConte (syn) sayi Blatchley (Chu, 1945; Emden, 1942) Anisodactylus binotatus Fabricius (Larsson, 1941) Anisodactylus californicus Dejean (Chu, 1945) Anisodactylus discoideus Dejean (Dogger and Olson, 1966) Anisodactylus sanctaecrucis Fabricius (Kirk, 1976) Geopinus incrassatus Dejean (Kirk, 1972) Amphasia interstitialis Say (Chu, 1945) Cratacanthus dubius Beauvois (Chu, 1945; Emden, 1942) Euryderus grossus Say (Chu, 1945) Harpalus affinis Schrank (syn) aeneus Fabricius (Schiodte, 1867, 1972) (syn) viridiaeneus (Chu, 1945) Harpalus bicolor Fabricius (syn) compar LeConte (Chu, 1945) Harpalus caliginosus Fabricius (Massey, 1893; Chu, 1945; Kirk, 1972) Harpalus erraticus Say (Kirk, 1972) | Harpalus erythropus Dejean (Chu, 1945) | Harpalus herbivagus Say (Lugger, 1899; Kirk, 1972) | Harpalus longicollis LeConte | (syn) vagans LeConte (Chu, 1945) | Harpalus pennsylvuanicus Degeer (Chu, 1945; Kirk, 1972) . Harpalus puncticeps Stephens (syn) anfusticollis Mueller (Larsson, 1941) Harpalus rufibarbis Fabricius (syn) seladon Schauberger (Larsson, 1941) Harpalus rufipes Degeer (syn) pubescens Mueller (Larsson, 1941) (syn) ruficornis Fabricius (Schiodte, 1867, 1872) Trichotichnus dichrous Dejean (Chu, 1945) Tribe LICHININI Dicaelus purpuratus Bonelli (Dimmock and Knab, 1904) (sbsp) splendidus Say (Wickham, 1893a ) Dicaelus sculptilis Say (Kirk, 1972) Licinus silphoides Rossi (Raynaud, 1970) Tribe CHLAENTINI Chlaenius cumatilis LeConte (Chu, 1945) Chlaenius impunctifrons Say (Claassen, 1919) Chlaenius laticollis Say (Schaupp, 1880a ) Chlaenius leucoscelis Chevrolat (Schaupp, 1880b ) VOLUME 90, NUMBER 1 105 Chlaenius pennsylvanicus Say (Chu, 1945) Chlaenius platyderus Chaudoir ( Kirk, 1972) Chlaenius prasinus Dejean (Chu, 1945; Schaupp, 1880b ) Chlaenius sericeus Forster (Wickham, 1895; Chu, 1945; Kirk, 1972) Chlaenius tomentosus Say (Kirk, 1972) Chlaenius tricolor Dejean (Chu, 1945) Tribe LEBIINI Callida viridicollis LeConte (Larson, 1969) Technophilus croceicollis Menetries ( Larson, 1969) (sbsp) peigani Larson (Larson, 1969) Plochionus timidus Haldeman (Larson, 1969; Duffey, 1891) Plochionus pallens Fabricius (Duffey, 1891) Lebia grandis Hentz (Chaboussou, 1939) Tribe GALERITINI Galerita janus Fabricius (Hubbard, 1875) Galerita lecontei Dejean (Salle, 1849) Tribe BRACHININI Brachinus cyanipennis Say (Clausen, 1940) Brachinus janthinipennis Dejean (Wickham, 1893b; Dimmock and Knab, 1904 ) Brachinus pallidus Erwin (Erwin, 1967 ) Acknowledgments I especially thank Henri Goulet for his comments and suggestions on the limits of this list prior to its compilation. I also thank Robert T. Allen for reading the manuscript and offering helpful suggestions and criticisms and Russ Ann Shannon for typing the manuscript. This study partially supported by USDA Cooperative Agreement No. 12-14-100-10, 644(33). Published with approval of the Director, Arkansas Agricultural Experiment Station, Fayetteville, Arkansas. Literature Cited Andersen, Johan. 1968. The larva of Miscodera arctica Paykull (Col. Carabidae). Norsk. Ent. Tidsskr. 15:71-74. 1970. The larvae of Pelophila borealis Payk., Nebria gyllenhali Schnh. and N. nivalis Payk. (Coleoptera, Carabidae). Astarte 3:87-95. Boldori, L. 1931. Nuovi appunti sulle larve dei Trechini. Le Grotte d'Ital. 5. 1932. Altri appunti sulle larve dei Trechini. Mem. Soc. Ent. Ital. 10, 106 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Boving, A. G. 1910, 1911. Nye Bidrag til Carabernes Udviklingshistorie I. II. Ent. Meddel. (N.S.) 3:319-376; 4:129-180. , and Craighead, F. C. 1931. An illustrated synopsis of the principal larval forms of the Order Coleoptera. Ent. Amer. (N.S.) 11l:pp. I-VIII, 1-351. Bryson, H. R., and Dillon, G. F. 1941. Observations on the morphology of the Corn Seed Beetle (Agonoderus pallipes Fab., Carabidae). Ann. Ent. Soc. Amer. 34(1):43-50. Burgess, A. F., and Collins, C. W. 1917. The genus Calosoma. Bull. U.S. Dept. Agric. ANT :1—124. Cerruti, M. 1939. Larva del Tachys parvulus Dejean, (Coleopt. Carab.). Mem. Soc. joie, Nall, Ih¢/e Ao Chaboussou, F. 1939. Contribution a letude biologique de Lebia grandis Hentz., predateur du Doryphore. Ann. Epipht. and Phytogen. (N.S.) 5:387-433. Chu, H.-F. 1945. The larvae of the Harpalinae Unisetosae (Coleoptera, Carabidae). Ent. Amer. 25(1):1-71. Claassen, P. W. 1919. Life history and biological notes on Chlaenius impunctifrons Say. (Coleoptera, Carabidae). Ann. Ent. Soc. Amer. 12:95-101. Clausen, C. P. 1940. Entomophagous Insects. McGraw-Hill Co., N.Y., pp. 528-533. Dimmock, G., and Knab, F. 1904. Early stages of Carabidae. Bull. Springfield Mus. Nat. Hist. 1:1—56. Dogger, J. R., and Olson, C. A. 1966. Larval characteristics of some North Dakota Carabids (Coleoptera: Carabidae). Coleopt. Bull. 20:91-96. Duffey, J. C. 1891. Transformations of a Carabid (Plochionus timidus), and c. Trans. Acad. Sci. St. Louis 5:533-542. Emden, F. I. van. 1942. | an Bis 154 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 4. Klausewitzia aphanes, new species, USNM 213783, 14.9 mm SL; locality un- known. Lateral view of premaxillary and maxillary bones. Anterior is to right. rather blunt snout. Profile along root of dorsal fin slightly convex. Body profile posterior to dorsal-fin origin concave to end of body. Ventral body profile nearly straight (in males) to gently convex in females with eggs. Entire dorsal fin anterior to vertical line through anus. Pelvic-fin origin posterior to vertical line through anterior origin of dorsal fin. Anterior anal-fin origin posterior to vertical line from posterior dorsal-fin origin. Caudal peduncle long, slender. Head large. Snout moderately blunt. Mouth terminal. Eye large. Bony interorbital moderate. Teeth all simple, conic. Two rows of teeth on dentary, anterior (outer) row with [8] (7-8) teeth, inner row [11] (11-13) teeth; premaxillary with one row of [8] (8-9) teeth; maxillary with one row of [9] (9-17) teeth; ectopterygoid with one row of (6-8) teeth in two alizarin preparations. Branchiostegal rays 3 in two alizarin preparations; ceratohyal with 2 rays, and 1 ray articulated between ceratohyal and epihyal. Gill rakers 4+ 8 in two alizarin preparations. Frontal-parietal fontanel of moderate size, completely separating parietal bones and separating frontals only at their posterior median margins. In two alizarin preparations first (anterior ) circumorbital bone ossified, with a laterosensory canal and second orbital bone ossified anteriorly and with a canal in that region. Antorbital and supraorbital bones not ossified, apparently absent in two alizarin speci- mens. Scales cycloid with up to 5-6 radii on exposed field in two alizarin preparations, lateral line incomplete, of [4] (4-5) perforated scales. Scales in a lateral series [33] (31-33), usually 32. Scales rows between dorsal and VOLUME 90, NUMBER 1 15 Ol Fig. 5. Klausewitzia aphanes, new species, USNM 213783, 14.9 mm SL; locality unknown. Lateral view of dentary bone. Note outer (anterior) tooth row lies on a more ventrally placed ridge of bone than inner (posterior) row. Anterior is to right. anal fin 8; 12 scale rows around caudal peduncle in all specimens. Pre- dorsal scales [9] (9-10). Area on belly anterior to pectoral fin bases with- out scales. Dorsal-fin rays ii,8,i in all specimens with locality data, some aquarium specimens with ii,8,i; ii,9; ii,9,i; or 1,10. Adipose fin present or absent, absent in holotype. Anal-fin rays [1i,6] (ii,6; ii,6,i; ii,7; or ii,7,i), usually ii,6 or Fig. 6. Klausewitzia aphanes, new species, USNM 213783, 14.9 mm SL; locality unknown. Medial view of dentary bone. Note 5 small replacement teeth shown posterior to inner (posterior) row of teeth. Anterior is to left. 156 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON ii,7. Pectoral fin inserted low on body, not flattened at base to provide a flat surface to substrate as in some species of Characidium; pectoral fin elongate, reaching to or beyond origin of pelvic fin. Pectoral-fin rays [viii] ( viii-ix ), usually viii. In aquarium specimens grown to 15 to 16.5 mm SL pectoral-fin rays became branched, having rays i,7 or i,6,i. Pelvic-fin rays reaching posteriorly beyond anus and anterior origin of anal fin in several specimens, especially males. Caudal fin forked, with 17 branched rays in all specimens; principal ray count 10/9 in all specimens. Total vertebrae including Weberian apparatus and terminal half centrum [33] (three paratypes with 32 and two with 33). Specimens from Lago José-Acu at Parintins have one with 32 and two with 33 vertebrae while the aquarium specimens have three with 31 vertebrae, six with 32 vertebrae and five with 33 vertebrae. First and second hypurals not fused, supra- neurals present, 3 or 4. At least 2 postcleithra on each side; cleithra an- teriorly articulate with each other in a specialized but movable joint. Coracoid of each side with their medial plates not in contact or parallel with each other, diverging laterally from their anterior point of contact. Color in alcohol.—Body color pale yellowish brown, nearly white, spotted profusely with small, apparently contracted brown chromatophores. These chromatophores oriented mostly along scale borders, especially on back, forming a reticulate pattern. A single median horizontal stripe beginning anteriorly at tip of snout and lower jaw, then passing posteriorly through maxillary bone and on to anterior border of eye. Posterior to eye horizontal stripe not obvious on head, but extending posteriorly across head just ventral to exposed portion of pterotic bone and continuing to region just dorsal to opercle where lateral-line scales begin. At this point horizontal stripe appears rather broad and merges with a dorsally elongate shoulder spot beginning at scale row just dorsal to lateral-line scale row and ex- tending ventrally to pectoral fin origin. Shoulder spot formed of moderately dense, contracted brown chromatophores. Shoulder spot of variable in- tensity, very dark in some males during life and in preservative. Horizontal stripe continues posterior to shoulder spot as a broad line to a point on an imaginary vertical line extended ventrally from anterior origin of dorsal fin. At this point, horizontal stripe becomes a single series of dark brown chromatophores situated along body at junction of epaxial and hypaxial muscle masses. Horizontal stripe continues posteriorly as a single row of chromatophores to a point on an imaginary vertical line extending dorsally from posterior termination of anal-fin rays. Here horizontal stripe becomes diffuse and lost among scattered brown chromatophores of caudal peduncle. Top of head dark brown, covered with chromatophores of moderate size. Up to eight transverse, brown dorsal saddle marks across back. Saddle marks conspicuous because pigment of scale borders, which is much darker VOLUME 90, NUMBER 1 157 than that of scale centers, is formed of a broad band of dark brown chromatophores. Two of these saddle marks situated between nape and a third saddle mark which occurs just anterior to base of dorsal fin. Fourth saddle mark lies at base of midregion of dorsal fin and fifth saddle mark located at or just beyond posterior termination of dorsal fin. Three addi- tional saddle marks may occur between fifth saddle mark and posterior termination of caudal peduncle. Posterior to eighth saddle mark is a cen- tral dark spot on dorsal surface of posterior portion of caudal peduncle. A ventral saddlelike mark also occurs at anterior and posterior base of anal fin and a third occurs on ventral surface of caudal peduncle near its pos- terior termination. Sometimes sides of body in area ventral to narrow por- tion of longitudinal stripe with one or two narrow vertical bars. Intensity of saddle marks variable, sometimes very dark, sometimes very pale both during life and in preservative. Vertical length of saddle marks quite vari- able on live and preserved specimens. Several specimens with a small lateral spot on caudal peduncle at base of tail fin. All fins mostly hyaline but dorsal, anal, and pelvic fins with considerable amount of brown pigment along fin rays and fin membranes. Area of dorsal fin formed by anterior 3 rays quite dusky. At about one-third of distance of each ray length from body, all dorsal-fin rays have additional dark brown pigment. These pigment spots form a dusky stripe on fin parallel to profile of back. Color in life—Color much as in preserved specimens; no bright colors. Dark brown pigment of preserved specimens black in life. Body muscula- ture rather translucent, that of back and caudal peduncle olive green in color. Peritoneum bright silvery except posteriorly in females with well- developed eggs. Eggs pale yellow green in life and easily seen through body wall. Head white ventral to eye. Fins hyaline except for dark pig- ment noted in preserved specimens. Etymology.—The name aphanes is from the Greek meaning obscure or unseen and is in reference to the cryptic qualities of live specimens of this fish. Relationships—We place K. aphanes in the previously monotypic genus Klausewitzia strictly on the basis of typology because it, like K. ritae Gery (1965) bears maxillary teeth. K. aphanes lacks the specialization of ventral mouth and elongate head and body present in Ammocryptocharax, the other genus in the Characidiinae having maxillary teeth. See Weitzman and Kanazawa (1976) for discussion of Klausewitzia and Ammocrypto- charax. The genera of the Characidiinae are in need of revision from a phylogenetic point of view, a project we have in progress. We do not necessarily believe that K. aphanes is phylogenetically closely related to K. ritae, especially since the primary typological character “relating” them PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 158 "'8=N 4 ZN x Lib [rey i G8 8°8 —0'8 V8 66 —0'8 66 Yisue] yous OL hte) 18) fe 08 08 —0'8 o8 #28 —0'8 8 eae te eT © OL 2 COA (6) SCL O'SCI-G TI GIT 0'CI-0'°0T Sele Joyourvip aA CGE SVvE-8 TE oge O'9E-V'VE CVE t 9S-9'GE Tet yisusl peoH] O'VG L LO-&'€G L'0€ CTE-V'0E 9° LG 6'0€-9'SG G6 9G Yasue] Ulf-OlAlod 8°61 46'GE-8'8T 8°06 806-906 606 4V'CC-C'81 = YSUs, UlF-[e10}0q 61 VIG-98I © 61 0'06-G LT 9°81 66I-GAT 661 yysus, epunped jepneD 86 LOlSss 66 V'0I-9'6 66 S'O1-8'8 0'OT ywdep epunped [epneD TS CLL-LGL c'92 8'9L-0'9L VSL 0'08-0'0L 691 USO Ulf-[eUe 0} JoUsS Des O'VS—L 6h Ore Oe Site) US VCS 9'7S-0'8P SEG UISTO UTF-OFATod 0} ynousg 9'SP 8'0S-T OV 80S © WIL OS 00S & 6S 8 GP €'SGg UIBHO UlF-][eSIOp 0} yous G'9G G86-L VG & GG 9'SS-8'VG CVG V 96-G'EG VSG ydop 4sozvo.lr) Ii SOI SUSI Lol (SITS) ll CGI OET-OTTI OST Yisus] prepurys x (6=N) x (€=N) x (L=N) 9dAjojoH ISULY ISU ospUuRy OLSON OLY WOLF ATGeqo.id ‘suouloods wmntirenby SUIJULIV | “SBUOZRULY OT vavnonandR T, ‘O1BON OL] ‘soupydp pizyMasnyyy JO soLQowmoydioyy "T a[qey, VOLUME 90, NUMBER 1 159 is the presence of maxillary teeth, a character probably primitive for the subfamily. See Weitzman and Kanazawa (1976) for a brief discussion of the primitive nature of maxillary teeth. The original definition of Klausewitzia by Géry (1965) needs much revision if K. aphanes is to be included. For example, in addition to the presence of maxillary teeth, Klausewitzia was originally proposed and defined by having the lateral line complete (it is incomplete in K. aphanes), an adipose fin present (it is present or absent in K. aphanes), the isthmus covered with scales (they are absent in K. aphanes), with the pectoral-fin rays not branched (they are branched in the larger specimens and unbranched in the smaller speci- mens of K. aphanes), fontanels absent (they are present in K. aphanes), and with both bicuspid and conical maxillary and premaxillary teeth (they are all unicuspid in K. aphanes). We do not think that many of these characters can be used as a satisfactory test of phylogenetic relationships because of their possible lability and because several of them are reduc- tive and perhaps convergent. See Weitzman and Fink (in press) for dis- cussion of the problem of convergence in small characoid fishes. Further- more some of these characters are labile in K. aphanes, for example the presence or absence of the adipose fin. These are certainly not char- acters indicating stability and therefore we suggest they may possess a very low degree of phylogenetic information for consideration of relation- ships at the generic level. They are not characters suitable for testing re- lationship in these fishes and we therefore reject them as characters for the description of genera in the Characidiinae. Literature Cited Bailey, Reeve M., et al. 1970. A list of common and scientific names of fishes from the United States and Canada (third edition). Amer. Fish. Soc. Special Pub. no. 6, Washington, D.C. 150 pp. Branson, Branley A., and G. A. Moore. 1962. The lateralis components of the acoustico-lateralis system in the sunfish family Centrarchidae. Copeia 1962(1): 1-108. Géry, J. 1965. Poissons characoides sud-americains du Senckenberg Muséum, II. Characidae et Crenuchidae de l’Igarapé Préto (Haute Amazonia). Senck. Biol. — 46(3):195-218. Weitzman, Stanley H., and W. L. Fink. (in press). Interrelationships of the neon tetras, a unique group of South American characids (Pisces), with comments on the interpretation of the phylogeny of New World characoid fishes. Smith- sonian Contrib. to Zool. Weitzman, Stanley H., and R. H. Kanazawa. 1976. Ammocryptocharax elegans, a new genus and species of riffle inhabiting characoid fish (Teleostei: Characidae ) from South America. Proc. Biol. Soc. Washington 89( 26) :325—346. Department of Vertebrate Zoology, National Museum of Natural History, Washington, D.C. 20560. 160 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Footnote *We follow Bailey, et al. (1970) in placing Elassoma in the Centrarchidae. Branson and Moore (1962) maintained the family Elassomidae for the genus Elassoma. They hypothesized that their Elassomidae was not related to the Centrarchidae. However, the many characters they listed on p. 76 separating the two families, such as reduced size of orbital bones, and reduced ossification of head canal bones and reduction in associated soft structures are common character states associated with pygmy fishes in many teleost groups such as small gobies, characoids, and cyprinids. We do not find that Branson and Moore demonstrated any fundamental differences between the Centrarchidae and their Elassomidae and therefore do not accept family status for the pygmy sunfishes. PROC. BIOL. SOC. WASH. 90(1), pp. 161-171 A NEW MARINE GENUS AND SPECIES OF THE NUUANU- GROUP (CRUSTACEA, AMPHIPODA) FROM THE YUCATAN PENINSULA Larry D. McKinney and J. Laurens Barnard Our colleague, Dr. E. L. Bousfield, National Museum of Canada, is in the process of dividing the Gammaridae (sensu lato) into many new families and superfamilies. We believe that the 4 genera, Gammarella Bate, 1857 (= Pherusa Leach, homonym Polychaeta, = Pherusana J. L. Barnard, 1964), Nuuanu J. L. Barnard, 1970, Cottesloe J. L. Barnard, 1974, and Tabatzius, new genus (described below), comprise a group distinguished by the miniaturized uropod 3, the serrate ventral margin on article 2 of pereopods 5-7, the unusual shield-shaped article 2 of pereopod 7, the form of teeth and spines on urosomites 2-3, and the oddly geniculate antenna 1. The Nuuanu-Group of Genera Diagnosis.—Uropod 3 miniaturized, of parviramus form but shorter than uropod 2, inner ramus shortened and scalelike, outer ramus in primitive genera greatly longer than inner ramus and bearing distinct article 2, but in advanced genera also becoming reduced and article 2 becoming vestigial or Pabsent. Antenna 1 geniculate between articles 1-2, occasionally be- tween articles 2-3. Article 2 of pereopod 7 much broader than on pereo- pods 5-6, bearing either large posterior serrations or small ventral serra- tions, or of expanded nasiform shape. Simple gills present on coxae 2-6. Lower lip lacking inner lobes. Inner plates of maxillae 1-2 densely setose medially. Gnathopod 2 dimorphic; male gnathopod 2 enlarged, article 6 very large, longer than article 5, palm and posterior margin confluent, bearing dense medial fuzz or tufts of elongate setae, dactyl curving to fit posterior margin; female gnathopod 2 small; gnathopod 1 of both sexes with medial fuzz or scales on articles 4-5; female gnathopod 2 with fuzz or scales medially on articles 2-6. Pleon dorsally smooth or toothed, urosomites 2-3 each bearing dorsolateral tooth on each side with attendant spine. Uropod 1 with basofacial spine on peduncle. Etymology.—A familial name characterizing this group may be taken from the second described genus in the complex owing to the similarity between Gammarella Bate, 1857, and Gammarellus Herbst, 1793. The latter name undoubtedly will be used as root for a new name of the family-group in revisions being undertaken by our colleagues. Both names are valid according to ICZN 56a. Relationship.—Taxa of the Nuuanu-group have the thick-bodied appear- 162 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON ance of those in the Elasmopus-group but differ strikingly in uropod 3 which is of melitid form, basically parviramus but miniaturized, and more primitive than that found in many species of Melita Leach owing to the well developed article 2 on the outer ramus, characteristic of certain other genera such as Eriopisa Stebbing, Ceradocopsis Schellenberg (= Maera- cunha Stephensen) and Metaceradocoides Birstein and Vinogradova. The latter genus may actually be related to Bathyceradocus Pirlot, which must be segregated from other marine amphipods because of the coxal gill on pereonite 7, and which may have affinities with another marine group typified by Anisogammarus Derzhavin. The lack of inner lobes on the lower lip of the Nuuanu-group distinguishes it from most other marine amphipods lacking gill 7 except for the Hadziidae (see Zimmerman and Barnard, 1977), a family composed of Hadzia Karaman, Dulzura J. L. Barnard, and Protohadzia Zimmerman and Barnard (1977). The Hadziidae have been distinguished from the Melitidae by Zimmer- man and Barnard (1977) by loss of the palm on female gnathopod 2. The weckeliid genera have been removed from the Hadziidae and differ- entiated from both Melitidae and Hadziidae by the enfeebled gnathopod 2 bearing evenly distributed spines on the palm. Weckeliids differ from Hadziidae also in the aequiramus uropod 3 lacking article 2 on the outer ramus (see Zimmerman and Barnard, 1977, for terms). One might conceive that Dulzura J. L. Barnard, in the Hadziidae, and Cottesloe J. L. Barnard in the Nuuanu-group could have had a common ancestor. In other respects the Nuwanu-group may be closer to hadziids than is apparent from the gross external appearance. Hadziids are an incipient interstitial group with thin bodies, no eyes and no pigment (except Protohadzia Zimmerman and Barnard, 1977). The Nuuanu-group have stout, Elasmopus-like bodies with stout appendages, but are probably almost pigmentless and the eyes in most species have few ommatidia. One species has been found in dimly lit anchialine waters of Hawaii (Bar- nard, in press). Both groups are therefore preadapted for lightless environ- ments, though neither has been found in the deep sea and both occur in shallow water when marine. The Nuuanu-group is very close to Ceradocopsis Schellenberg (= Maeracunha Stephensen) which also has miniaturized uropod 3, similar lower lip, and similar gnathopods, but the Nuuanu-group has article 2 of pereopod 7 broadly expanded and hatchet shaped. The retention of medial setae on the maxillae distinguishes the Nuuanu- group from the Elasmopus-group of genera, which, broadly conceived, would include Parelasmopus Chevreux, Mallacoota J. L. Barnard, Ifalukia J. L. Barnard and Beaudettia J. L. Barmmard. A tendency to reduction of the inner ramus of uropod 3 occurs in Elasmopus, and this is carried to the parviramus extreme in the aberrant Beaudettia. Elasmopus- and Nuuanu- VOLUME 90, NUMBER 1 163 groups might conceivably have a common ancestor in the morphological grade represented by Ceradocopsis. The Eriopisella-group has reduced maxillary setation and severely re- duced gnathopod 2 approaching the classification “mitten-shaped.” The separation of the Nuuwanu-group from other marine gammarids is therefore very strong and worthy of nomenclatural recognition. Classification in the Nuuanu-group An opportunity to examine one specimen of Gammarella fucicola (Leach), see below, reveals the extreme closeness of Gammarella and Cottesloe J. L. Barnard, 1974. In the following key an attempt to distinguish them is made, but the diversity between the 2 known Australian species of Cottesloe is greater than between the type-species of Cottesloe, C. berringar, and Gammarella fucicola. There may be reason to separate the second species of Cottesloe, C. merringannee, into a third genus or simply to combine all into an amended Gammarella. This latitude could be extended to admit Nuuanu into the expanded genus also. Until more exploration of warm waters is completed and more species of the complex discovered we maintain the genera in the following key. Key to Genera and some Species of the Nuuwanu-group 1. Maxilla 1 styliform T abatzius Maxilla 1 not styliform 2 2. Urosomite 1 with dorsal tooth or process 3 Urosomite 1 smooth dorsally 4 3. Article 2 of antenna 1] about 1.25 times as long as article 4 of antenna 2, cuticle covered with straw setules Gammarella fucicola Article 2 of antenna 1 about 1.05 times as long as article 4 of antenna 2, cuticle naked Cottesloe berringar 4, Article 2 of antenna 1 about 0.8 times as long as article 1 of antenna 1, cuticle naked, pleonites 1-3 with dorsal tooth Cottesloe merringannee Article 2 of antenna 1 about 0.55 times as long as article 1 of antenna 1, cuticle covered with straw setules, pleonites 1-3 untoothed dorsally Nuuanu Tabatzius, new genus Diagnosis —Nuuanuid with styliform outer plate and palp of maxilla 1, article 1 of palp elongate; mandibular lobes of lower lip elongate, apically curled; lobes of telson with deep apical notch; head with mammilliform lateral lobe defined by weak notch; uropod 3 with outer ramus longer than peduncle, distinctly 2-articulate, inner ramus short and scalelike. 164 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Type-species—Tabatzius copillius, new species. Etymology.—Derived from Mayan words, “Tabatzi,’ morning star; “copilli,” wedge-shaped. Gender masculine. Relationship.—This genus is characterized by the remarkable styliform maxilla 1 and the elongate and curled mandibular lobes of the lower lip. There is a possibility that the species of Nuuanu, the most similar genus, will have to be segregated generically in the future but Tabatzius can be distinguished from the 3 known species of Nuwanu by the first maxilla and lower lip. No other member of the Nuuanu-group has deeply notched telsonic lobes, although several species such as Cottesloe berringar have weakly excavate apical margins. The head in this group varies in the angularity or roundness of the ocular lobe and this may have future value at the generic level. We suspect that the 2 species of Cottesloe may be distinct generically owing to differences in epimera 1-2, teeth on article 2 of pereopods 5-7, and teeth on pleonites. Gammarella, as depicted by Della Valle (1893), and Chevreux and Fage (1925), has normal mouthparts but very weakly toothed second articles on pereopods 5-7 and an unusual male uropod 3 with reduced article 2 on the outer ramus. It must be reexamined to determine whether or not uropod 3 is similar to that of Cottesloe berringar (see below). Gnathopod 2 of the male also has an unusually long dactyl and an unusual cluster of basal setae on article 6. Tabatzius has an unusual female gnathopod 2. It is larger and more distinct from gnathopod 1 than in females of other species in the family. Tabatzius copillius, new species Figs. 1-3 Description of male.—Lateral lobe of head mammilliform, anteroventral notch weak; eyes with numerous but dispersed ommatidia, all circular, none bigeminal. Article 1 of antenna 1 with distoventral spine, accessory flagel- lum 3-articulate, primary flagellum with 11 articles, as long as peduncle. Articles 4 and 5 of peduncle on antenna 2 of equal length; flagellum with 6 articles, about 1.4 times as long as article 5 of peduncle. Right lacinia mobilis with 3 prongs, one of these short; right rakers 8, left rakers 8, one setule between each pair of rakers; palp rathe: short and stout, ratio of lengths of articles 1,2,3 = 27:53:44, article 2 with only 1 stout inner apical spine, article 3 scarcely falcate, inner apical spines inserted basally to mark 45. Lower lip elongate proximally to distally; mandibular lobes strongly extended, thin, curled apically, with ridge ending in lobe on each side proximal to curled lobe of tip. Inner plate of maxilla 1 thin, with mediofacial setose ridge and apical extension free of setae; outer plate and palp styliform, extremely elongate, thin; outer plate with 3 stout apical VOLUME 90, NUMBER 1 165 Xie Fig. 1. Tabatzius copillius, new genus, new species, holotype, male “a”; >» = male “b”; c = female “c.” A, antenna; C, coxa; D, dactyl of pereopod; E, epimeron(a); F, accessory flagellum; G, gnathopod; H, head; I, inner ramus or plate; J, palp; K, cuticle; L, lower lip; M, mandible; N, right lacinia mobilis; O, outer plate or ramus; P, pereopod (3-7); R, uropod; S, maxilliped; T, telson; U, upper lip; W, pleon; X, maxilla; e, broken or absent; f, flattened; i, medial; n, right; 0, opposite; s, setae removed; v, ventral. 166 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 2. Tabatzius copillius, new genus, new species, holotype, male “a.” See Fig. 1 for legend. spines, medialmost with 2 inner humps and notches, 2 additional free apical spines, 6 other thin inner facial spines, some dispersed towards base; palp 2-articulate, article 1 elongate, article 2 about 1.2 times as long as article 1, apex with tooth, bearing 4 apical spines, 1 of these pointing somewhat VOLUME 90, NUMBER 1 167 OR1 *. b = male Fig. 3. Tabatzius copillius, new genus, new species, holotype, male “a’ “b’; c = female “c.” See Fig. 1 for legend. laterally, plus 1 stiff subapical seta. Mavxilla 2 very thin; outer plate much narrower than inner plate, almost rectangular in apical part. Coxae 1-3 with small posterodistal tooth, coxae 1-2 with 2 posterior spines, coxa 3 with 1 posterior spine, posteroventral lobe of coxa 4 almost fully rounded. 168 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Long posterior setae on article 2 of pereopods 1-4 = 2-1-1-1, short posteriors = 1-0-4-2, long anteriors = 0-0-1-1, short anteriors = 2-0-3-1. Palm of gnathopod 1 oblique; article 2 expanding distally; article 3 not elongate; article 4 with 1 long and 6 medium to short posterior spines from lateral view, with medial fuzz and additional posterior spines; article 6 with only 1 anterior subapical seta, mediofacial setal groups highly posterior. Tooth on article 4 of gnathopod 2 strong, thin, pointing distalwards; posterior margin of article 6 with 3 notches each bearing setule but otherwise lacking long setae, fuzz on mediofacial hollow very sparse, anterofacial margin with only 3 short setae in tandem, distal part of ridge defining hollow with hump, and 2 spines distalwards from hump, lateral palmar margin with only 5 spines in groups of 3,1,1. Pereopods 3-4 alike; main dactylar seta almost fully fused to body of dactyl, posterior margin other- wise naked; coxa 5 with distomedial spine. Pereopods 5-7 short; posterior margins of article 2 fully toothed, teeth enlarged. No pereonite or pleonite with dorsal tooth; epimeron 1 with small posteroventral tooth and 2 antero- facial spines; epimeron 2 strongly extended posteroventrally, with sharp but stout posteroventral extension, ventral face with 4 spines; epimeron 3 with small to medium posteroventral tooth, anteroventral corner with spine. Urosomites 2-3 with dorsolateral tooth bearing spine on each side. Peduncle of uropod 1 with 1 small apicodorsal spine and special enlarged spine set more ventrally, dorsal margin with 2 spines set proximally, dorso- medial margin with 1 large special spine and 2 smaller dorsal spines set apicalwards; outer and inner rami with 2 dorsal spines. All rami of uropods 1-2 with 5 apical spines, comprising 1 large apical nail, 2 large accessory spines, and 2 small striate falcate spines, 1 of these on each side of apical nail, pair of large accessory spines on uropod 2 of diverse sizes. Peduncle of uropod | with 1 subapical dorsolateral spine, medial apex with 1 small spine; outer ramus with 2 dorsolateral spines, 1 dorsomedial spine; inner ramus with 3 dorsal spines. Inner ramus of uropod 3 small and scalelike, naked; outer ramus 1.2 times as long as peduncle, article 1 of outer ramus with 1 lateral spine. Telson somewhat elongate, ratio of width to length 27:31; apices narrow, deeply bifid, each with elongate spine inserted into notch plus short adjacent lateral setule, pair of dorsolateral setules on each side near mark 60. Cuticle covered densely with pores, often pores with attached short spicules but most spicules apparently abraded and absent. Description of female —Gnathopod 2 distinctive, thin but not like gnatho- pod 1; article 4 lacking posterodistal tooth, bearing brush of distal setae, naked on posterior margin; article 5 expanded, especially broad distally, posterior margin strongly bulging, anterior margin with 1 spine sub- apically; article 6 as long as article 5, somewhat more ovate than in other species of family, palm oblique but defined by precise angle, bearing pair of defining spines, 1 on each face, medial surfaces of articles with patches VOLUME 90, NUMBER 1 169 of ornamentation, either fuzz composed of elongate scales, or tiny buttons appearing as truncate villi, as follows: article 2 with distal scale fuzz, articles 3 and 4 with villose fuzz, article 5 with anterior and posterior patches of scale fuzz, article 6 with posteroproximal patch of scale fuzz. Variations —Three males and 1 female known. Female larger and better developed than males, with more numerous spines on pereopods 5-7, generally more serrations on article 2 of pereopods 5-7, ridge on coxa 5 half as long as in male. Primary flagellum of antenna 1 with 14 articles; flagel- lum of antenna 2 with 10 articles; facial clump on article 1 of antenna 1 reduced to 4 setules. Maxillae more setose than in males. Inner ramus of uropod | with 3 dorsal spines; outer ramus of uropod 2 with 3 dorsal spines, inner with 4; lateral apex of peduncle on uropod 3 with 4 spines. Male “b” better developed than holotype male. Primary flagellum of antenna | with 11 articles (broken in holotype). Coxa 1 with 3 posterior spines, posterior margin of article 5 on gnathopod 1 with 7 rows of spines, coxa 5 with 3 posterior setules and slight notch (absent on holotype), article 2 of pereopod 6 with 18 serrations but article 2 of pereopod 7 with only 22 serrations. Outer ramus of uropod 1 with 3 dorsal spines. Notes on illustrations ——Eyes not drawn on head of holotype, see female head for eyes. Apex of antenna 1 broken (see description of male “b” for proportions ). Left pereopod 7 with article 6 stunted (see enlarged drawing of male “b”). Epimera of pleon drawing unflattened (see additional draw- ing of flattened epimera). Telson with several missing setules marked with sockets. Holotype —USNM 154430, male “a” 3.22 mm (illustrated). Type-locality—Smithsonian-Bredin 1960 Expedition, Sta. 52-60: 10 April 1960, Ascension Bay, Quintana Roo, Mexico, just behind center of Nicche- habin Reef, E of Allen: Point, 1-5 ft, on coral and under coral pieces on bottom. Paratypes.—Female “c” 3.68 mm (illustrated), from type-locality. Male “b” 3.30 mm (illustrated), and male “d” 2.30 mm, from Smithsonian-Bredin 1960 Expedition Sta. 95-60: 19 April 1960, Ascension Bay, Suliman Point to 300 yds to SW shore and reef flats to sand flats, 5 ft. Gammarella fucicola (Leach) Pherusa fucicola Leach, 1814:432; Stebbing, 1906:449-450; Chevreux and Fage, 1925:247-248, figs. 258, 259. Melita fucicola—Sovinsky, 1898:486-489, pl. 11, figs. 10-19, pl. 12, figs. 1-4. Material—USNM 172342, female 8.5 mm, Cap Coz, Finistere, Brittany, France, 23 August 1945, from seaweed, Andrew D. Pizzini. Remarks.—Except for Sovinsky’s fine portrayal of this species, the illus- trations and descriptions appear to be erroneous in several respects. The 170 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON characteristics of head, antenna | and uropod 3 fit the Nuwanu-Cottesloe- Tabatzius concept much closer than heretofore known. Although article 2 of antenna | is elongate it forms a geniculation with article 1, while article 3 also folds in an opposite manner as in the other genera. The head is similar to Nuuanu amikai J. L. Barnard, 1970, and not as shown by Chevreux and Fage (1925), who overlooked the notch and accessory lobe covering the base of antenna 2. Uropod 3 of our female is like that of N. amikai in the well developed article 2 of the outer ramus and in general proportions of the outer and inner rami, in contrast to the illustration of both Della Valle and Chevreux and Fage. Perhaps male uropod 3 changes to the form shown by Chevreux and Fage. The cuticle is covered with straw- setules; each side of urosomites 1-2 has a dorsal spine and quadrately ex- tended tooth as in other species of the family. Uropod 1 has a large fully apical spine on each side of the peduncle as in Cottesloe berringar. The medial pattern of fuzz-setules on male gnathopod 2 shown by Della Valle (1893) and Chevreux and Fage (1925) may be a generic distinction between Gammarella and Cottesloe. Acknowledgments We thank Biruta Akerbergs and Carolyn L. Cox for inking and lettering our plates. Charline M. Barnard undertook our research in the literature. Dr. J. H. Stock, Institut voor Taxonomische Zoslogie, Universiteit van Amsterdam, kindly offered us valuable information and advice. Literature Cited Bamard, J. L. 1964. Revision of some families, genera and species of gammaridean Amphipoda. Crustaceana 7:49-74, tables 1-2. —. 1970. Sublittoral Gammaridea (Amphipoda) of the Hawaiian Islands. Smith- sonian Contributions to Zoology 34:286 pp., 180 figs. 1974. Gammaridean Amphipoda of Australia, Part IJ. Smithsonian Con- tributions to Zoology 139:148 pp., 83 figs. ———. in press. The Cavernicolus fauna of Hawaiian lava tubes 9. Amphipoda (Crustacea) from brackish lava ponds. Pacific Insects 17. 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. Chevreux, E., and L. Fage. 1925. Amphipodes. Faune de France 9:488 pp., 438 figs. Della Valle, A. 1893. Gammarina del Golfo di Napoli. Fauna und Flora des Golfes von Neapel . . . Monographi 20:xi + 948 pp., atlas of 61 plates. Herbst, J. F. W. 1793. Garneelasseln. Onisci gammarelli. Part 6 in: vol. 2 of “Versuch einer Naturgeschichte der Krabben und Krebse nebst einer system- atischen Beschreibung ihrer verschiedenen Arten” :105-146 [pls. 34-36, not seen], Berlin und Stralsund. Leach, W. E. 1813/14. Crustaceology. Appendix. Edinburgh Encyclopaedia, 7: 429-437. Sovinsky, V. K. 1898. Vysshiia rakoobraznyia (Malacostraca) Bosfora, po materialam | VOLUME 90, NUMBER 1 171 sobrannym d-rom A. A. Ostroumovym v 1892 i 93 gg. Zapiski Kiev. Obshchestva 15:447-518, pls. 8-13. Stebbing, T. R. R. 1906. Amphipoda I. Gammaridea. Das Tierreich 21:1—806, figs. 1-127. Zimmerman, R. J., and J. L. Bamard. 1977. A new genus of primitive hadziid (Amphipoda) from Puerto Rico. Proceedings of the Biological Society of Washington 89(50):565-580, figs. 1-5. Biology Department, Texas A&M University, College Station, Texas 77843 and Department of Invertebrate Zoology, Smithsonian Institution, Washington, D.C. 20560. PROC. BIOL. SOC. WASH. 90(1), pp. 172-182 PENAEOPSIS EDUARDOI, A NEW SPECIES OF SHRIMP (CRUSTACEA: PENAEIDAE) FROM THE INDO-WEST PACIFIC Isabel Pérez Farfante During a revision of the genus Penaeopsis, I discovered that a very distinct Indo-West Pacific species had not been recognized previously, representa- tives having been repeatedly assigned to 2 other species, both named by Bate in 1881. As indicated in the list of material examined, my conclusions are based on a study of the Penaeopsis collected during the voyage of the Challenger, 1873-76 and identified by Bate, including the types of his species, “Penaeus rectacutus’ and “P. serratus.” Also examined were 6 specimens described by de Man, and the relatively large collection of Penaeopsis taken by the U.S. steamer Albatross during the Philippine Expedition, 1907-1909, which includes representatives of 3 of the 4 species found in the Indo-West Pacific. The method of measuring specimens and the terminology used below are described by Pérez Farfante (1969). The scales accompanying the illustrations are in millimeters. The materials used are in the collections of the British Museum (Natural History) (BMNH), National Museum of Natural History (USNM), and the Zoological Museum, Amsterdam (ZMA). Penaeopsis eduardoi, new species Figs. 14 Penaeus rectacutus.—Bate, 1888:266 [part], pl. 36, fig. 2z. —? Villaluz and Arriola, 1938:38, pl. 3 [not Penaeus rectacutus Bate, 1881]. Penaeus serratus.—Bate, 1888:268 [part], pl. 37, fig. 1”, lq [not Penaeus serratus Bate, 1881]. Metapenaeus rectacutus—Alcock and Anderson, 1894:145 [at least all males]. Parapenaeus rectacutus.—de Man, 1911:82.—de Man, 1913, pl. 8, fig. 26a—c. —Yokoya, 1933:9. Penaeopsis rectactus.——Kubo, 1949:321, fig. 1H; 8J; 19C; 23A-B; 36K-L; AT]; 58P: 76A, F; 78K; LISA-G; 119. Penaeopsis challengeri de Man, 1911: 76 [part, 2 from Siboga-Expedition sta 253|].—Ivanov and Hassan, 1976:4. Review of Literature The Indo-West Pacific members of the genus Penaeopsis often have been misidentified. Bate (1881) first gave brief diagnoses of 2 species under the names of Penaeus rectacutus and Penaeus serratus, and a few years later VOLUME 90, NUMBER 1 173 (1888) presented rather extensive descriptions of both, which have been responsible for much subsequent confusion. This confusion has been due mainly to the following: a. The posterior part of the telson is missing in the specimen which Bate (1888) designated the type of P. rectacutus (p. 268), as he stated and clearly indicated in the illustration of the entire animal (Pl. 36, Fig. 2). b. Bate (1888) assigned to P. serratus a male—the petasma of which he described and figured (PI. 37, Fig. 1’ )—that actually belongs to a different species, Penaeopsis eduardoi n. sp. c. The original descriptions and the drawing of the thelycum of P. serratus lack necessary detail, and the illustration of the telson is inaccurate. The material on which Bate (1881) based his diagnosis of Penaeus rectacutus was obtained in the Philippine Islands, the only locality cited by him. In 1888, however, he listed in addition to the type (mentioned on p. 268), 5 females caught off Matuku, Fiji Islands, at Challenger sta 173. He stated that “The telson in the typical specimen has two small articulating spines on each side, beyond which it has been broken off.” Referring to the 5 females he wrote “the form of the thelycum in these corresponds with that of the type [of P. rectacutus|, but differs from that of Penaeus serratus, with which they were found associated. In these specimens the telson is armed. with two teeth posterior to the lateral spines, and therefore Penaeus rectacutus may be only a variety [presumably of P. serratus].” I have ex- amined 3 of these 5 females, none of which belong to either species, but to the one described herein, as does the male Bate attributed to P. serratus. Fur- thermore, my examination of the telson of Bate’s specimen of P. serratus which was illustrated (1888, Pl. 37, Fig. 1z) as lacking movable spines, revealed the presence of 3 such spines on each side, instead of 2 as stated (Bate, 1888). The study of numerous specimens of Penaeopsis rectacuta obtained since 1888, leaves no doubt that, like “Penaeus serratus” (= Penaeopsis challengeri de Man, 1911), this species usually possesses 3 pairs of movable spines on the telson; only occasionally are there only 2 pairs, as was reported by Ramadan (1938). Thus, most specimens of Penaeopsis rectacuta ditter trom Penaeopsis balssi Ivanov and Hassan, 1976, and Penaeopsis eduardoi, n. sp.. the other 2 Indo-West Pacific species which exhibit only 2 pairs of movable spines. Also, the thelycum of P. rectacuta, although superficially resembling the thelyca of the other Indo-West Pacific Penaeopsis, exhibits various fea- tures (clearly indicated in the illustration presented by Bate, 1888, Pl. 36, Fig. 2”) by which the females may be recognized readily. Furthermore, the males of this species, which were not available to Bate, differ markedly from those of P. eduardoi. Following Bate’s erroneous assignation of the male in which the ventral costa of the petasma is produced distally into a long spinelike projection to “Penaeus serratus, afew authors relegated to this species not only the males, 174 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON but also the females of P. eduardoi. Others, apparently recognizing that the thelycum of the females accompanying the males of P. eduardoi was different from that of the females of “P. serratus,” resembling more closely that of P. rectacuta, assigned their specimens to the latter. This misidentifi- cation was made by de Man (1911; 1913) and Kubo (1949), both thus unknowingly having noted in their descriptive accounts, and represented in illustrations, features by which P. eduardoi may be distinguished from P. rectacuta. Diagnosis.—First rostral tooth situated opposite orbital margin. Telson bearing 2 pairs of movable spines, its terminal part hastate. Petasma with ventral costa produced into long distal spine projecting beyond row of cin- cinnuli. Thelycum with plate on sternite XIV bearing caudally pedunculate posteromedian protuberance; posteromedian projection of plate on sternite XIII conspicuously bifid caudally, and tooth on sternite XII broad basally, semiconical, elongate and directed anteriorly. Description.—Rostrum (Fig. 1) horizontal to somewhat upturned, straight or slightly sinuous (strongly convex in young); length proportionately in- creasing with size of shrimp, in adult relatively long, reaching at least mid- length of third antennular article and often overreaching peduncle (in large females ), its maximum length about 0.8 that of carapace. Rostral plus epigastric teeth 8-15 (increasing in number through juvenile stage), basal rostral teeth close together, ultimate 3 or 4 usually relatively widely spaced; first rostral tooth situated at level of orbital margin; epigastric tooth situated about 0.35 carapace length from orbital margin. Paired adrostral carinae low, sharp posteriorly, almost indistinct along anteriormost portion of rostrum, dorsal one running along bases of teeth. Postrostral carina high anteriorly, lower, although well defined, behind epigastric tooth ending at about posterior 0.4 length of carapace, and followed by small dorsal tubercle located near posterior margin of carapace. Antennal and hepatic spines well developed, subequal in size; pterygostomian spine conspicuous, antero- ventral angle of carapace broadly obtuse (Fig. 2a). Cervical carina elevated and sharp, accompanying sulcus well marked; its posterodorsal extremity located about 0.45 length of carapace from orbital margin, and relatively far from postrostral carina. Hepatic carina anteriorly high, sharp, sinuous, extending from below hepatic spine to apex of pterygostomian spine; hepatic sulcus well marked along carina, very shallow posteriorly. Branchiocardiac carina extending posterodorsally to near margin of carapace, indistinct in many large individuals. Antennular peduncle with length about 0.65 that of carapace, third article narrow and about 1.2 times as long as second; prosartema falling short of distal margin of first article; stylocerite ending in small spine, length about 0.4 that of first article; distolateral spine long, slender and sharp, reaching between basal 0.65 and distal margin of second article; parapenaeid spine VOLUME 90, NUMBER 1 ISITE 2 9939= 99993559929999:93999999920909992993IFIIINIINIIA DErPPB Es) PEEIAIATIIFZIIA IDI 10 aaarnnAs97aIIAVAR IANS = 182:99979199995995~ 4 Peep 4 op oN Fig. 1. Penaeopsis eduardoi, holotype @, 27 mm cl, Balayan Bay, Luzon I, The Philippines: Lateral view. very long, considerably overreaching distal margin of article. Lateral flagel- lum as long as, or longer than carapace, not tapering, broad proximally, filiform distally; ventral flagellum sexually dimorphic: in female straight, tapering to filiform terminal part, its length about 0.8 that of carapace; in male deeply concave basally forming semicircle, latter joining straight distal part by dorsally arched thickening. 176 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 2. Penaeopsis eduardoi, holotype: a, Enlargement of anteroventral part of carapace; b, Telson and right uropod. Scaphocerite extending to, or barely surpassing, antennular peduncle; lateral rib ending in sharp spine falling slightly short of distal margin of lamella. Antennal flagellum broken in specimens examined, but not less than twice body length: in female 136 mm total length, incomplete flagellum 268 mm long. Third maxilliped of male extending as far as distal 0.35 of third antennular article, that of female to distal margin; ratio of dactyl: propodus about 0.70 in male, and 0.75 in female. First pereopod reaching about distal end of carpocerite, armed with spines on basis and ischium. Second pereopod overreaching carpocerite by length of dactyl or by almost entire propodus (also reaching at least distal 0.4, at most 0.1, of first antennular article), with basis and ischium unarmed. Third pereopod of male reaching between proximal 0.35 and distal end of second article, that of female, between midlength and distal end of third article. Fourth pereopod extending to distal end of carpocerite or surpassing it by length of dactyl, thus about as far as first. Fifth pereopod reaching between base and midlength of second article. Order of above appendages in terms of their maximum anterior extensions: first, fourth, second, third and fifth pereopod, and third maxilliped. Abdomen with dorsal keel on fourth to sixth somites, posterodorsal margin of fourth and fifth with median incision; length of sixth somite about 1.7 times maximum height, bearing long, usually interrupted cicatrix on VOLUME 90, NUMBER 1 Wer 4 Fig. 3. Penaeopsis eduardoi, 6, 16.5 mm cl, off Matuku, Fiji Islands: a, Lateral view of left half of petasma. b, Ventral view of petasma slightly extended; c, Dorsal view of same petasma; d, Dorsal view of right appendix masculina. (Illustrations pre- pared from stained specimens. ) 178 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON lateral surface, conspicuous spine at posterior end of keel, and pair of minute spines posteroventrally. Telson (Fig. 2b) with median sulcus flanked an- teriorly by 2 pairs of slender ridges, lateral pair extending to fixed spines; latter very long, in young extending as far as apex of telson; lateral margins of telson armed with 2 pairs of small, slender, movable spines, anterior pair situated about 0.4 length of telson from anterior margin, posterior pair slightly closer to other movable spines than to fixed ones; terminal part hastate, its length 6 to 7 times basal width. Mesial ramus of uropod reach- ing, or slightly overreaching, apex of telson; lateral ramus surpassing mesial one by almost 0.2 of its own length. Petasma (Fig. 3a-c) with ventral costa marginally situated distomesially (where bent inward) and continuing along curved distomesial margin at angle of about 40° to shaft of petasma; distal part of costa free from, though closely appressed to, margin of dorsolateral lobule, forming long spine projecting anterodorsally (sometimes also toward median line) beyond row of cincinnuli (hooklike structures along mesial margin of median lobes of petasma that serve to interlock its 2 halves). Ventromedian lobule with proximal plate flush with surrounding membranous portion, lacking mesial crest. Appendix masculina (Fig. 3d) transversely oval, broader than long, strongly convex dorsally, and bearing short setae around entire margin. Thelycum (Fig. 4) with paired anterior borders of plate of sternite XIV strongly arched, sloping posterolaterally, and separated by emargination receiving posteromedian projection of sternite XIII; lateral borders con- stricted (sharply turning mesially behind midlength) before joining pos- terior ridge; plate conspicuously overlapping sternite XIII, densely setose anteriorly, strongly slanting dorsomesially toward deep anteromedian part, and armed with short, caudally pedunculate posteromedian protuberance. Median plate of sternite XIII semicircular to subcordiform (with blunt apex), flat, studded with setae, its posteromedian projection caudally bifurcate. Sternite XII with posteromedian, subconical, broad (rather than compressed) tooth, and pair of strong ribs across posterolateral borders. Maximum carapace length, males 26 mm; females 34 mm. Material examined.—Holotype: ¢?, USNM 168298, 27 mm carapace length; about 120 mm total length; 22 mm rostrum length; epigastric plus 11 rostral teeth. Type-locality: Balayan Bay, Luzon I, The Philippines, 13°41’/00’N, 120°47’/05’E, 366 m, Albatross sta 5116. Paratypes—9é 72, USNM 168299, collected with the holotype. Other specimens.—Fiji Islands. 16 32, BMNH, off Matuku, 576 m, 24 July 1874, Challenger sta 173. South China Sea. 12, USNM, SW of Taiwan, 491 m, 5 November 1908, Albatross sta 5317. The Philippines. Luzon: 14, USNM, off Hermana Menor I, 326 m, 22 November 1908, Albatross sta 0331; 12, USNM, Balayan Bay, 324 m, 17 January 1908, Albatross sta 5112; VOLUME 90, NUMBER 1 179 Fig. 4. Penaeopsis eduardoi, holotype: Thelycum. 1é, USNM, Albay Gulf, 368 m, 8 June 1909, Albatross sta 5459. Bohol Strait (between Cebu and Bohol): 12, USNM, 296 m, 23 March 1909, Albatross sta 5412; 3¢ 12, USNM, 402 m, 9 April 1908, Albatross sta 5198. Mindanao: 36 12, USNM, off Tagolo Point, 401 m, 20 August 1909, Albatross sta 5541; 1é 62, USNM, off Tagolo Point, 366 m, 9 August 1909, Albatross sta 5518; 12, USNM, Macalajar Bay, 366-402 m, 5 August 1909, Albatross sta 5504-5; 1é 12, USNM, off El Salvador, 391-413 m, 4 August 1909, Albatross sta 5002-3; 16, USNM, Iligan Bay, 494 m, 5 August 1909, Albatross sta. 5508. Indonesia. 3°, ZMA, Makassar Strait, 450 m, 8 June 1899, Siboga sta 74. 12, ZMA, off Kai I, 304 m, 10 December 1899, Siboga sta 253. 16, ZMA, Bali Sea, 289 m, 14 March 1899, Siboga sta 12. 1°, ZMA, otf Paternoster Is, 521 m, 1 April 1899, Siboga sta 38. Comparative specimens of other species: Holotype 2 Penaeus rectacutus 180 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Bate, 1881, BMNH, between Bohol and Cebu, The Philippines, 95 fm (174 m), 22 January 1875, Challenger sta 209. Syntypes 42 Penaeus ser- ratus Bate, 1881, BMNH, off Matuku, Fiji Islands, 315 fm (576 m), 24 July 1874, Challenger sta 173. Geographic and bathymetric ranges.—This species has been found off the Fiji Islands and from Japan through The Philippines and Indonesia to the southwestern part of the Bay of Bengal, in depths between 290 and 970 m. Etymology.—This species is named for my son, Eduardo Canet. Discussion.—As stated above, Penaeopsis eduardoi differs from both P. rectacuta and P. challengeri de Man, 1911 [replacement name for Penaeop- sis serrata (Bate, 1881)], in possessing 2 pairs of movable spines on the telson, a character it shares with P. balssi, as well as with the amphi-Atlantic P. serrata Bate, 1881, which only rarely bears 3 pairs (Burkenroad, 1934). Penaeopsis eduardoi, however, can be separated from all its 4 congeners by the external genitalia. It is the only species in which the ventral costa of the petasma is produced into a long distal spine projecting beyond the row of cincinnuli. The thelycum, in turn, is unique in that the plate on sternite XIV exhibits a caudally pedunculate posteromedian protuberance, whereas in all the other species the latter is represented by a ridge or protuberance which is broad caudally or lacks a peduncle. Also the posteromedian projection of the plate on sternite XIII is conspicuously bifid caudally whereas in the other species it is straight or broadly emarginate, except in occasional females of P. rectacuta in which a shallow incision is present. Furthermore, sternite XII is armed with an elongate, broad basally, semi- conical, anteriorly directed tooth, which in the other members of Penaeopsis is absent or, if present, either laterally compressed, or short, conical and directed ventrally. Because the thelycum of P. eduardoi has been considered identical with, or very close to that of P. rectacuta, the differences between the two are given. In P. eduardoi the anterior borders of the plate of sternite XIV are directed posterolaterally and the lateral ones constricted, whereas in P. rectacuta the anterior borders are subhorizontal, the lateral ones virtually straight. Also in the latter species the protuberance born on the plate of XIV is caudally broad (often continued anteriorly as a ridge), the median plate of sternite XIII is cordiform (sharply pointed anteriorly) and con- cave, and the tooth on sternite XII is strongly compressed. In addition to the differences in the external genitalia as well as in the number of telsonic spines pointed out above, P. eduardoi may be distinguished from P. rectacuta by the position of the first rostral tooth, situated opposite the orbital margin in the former whereas conspicuously posterior to it in the latter. My examination of the specimens collected during the Siboga Expedition and studied by de Man (1911), demonstrated that the juvenile female VOLUME 90, NUMBER 1 181 from sta 253, which he identified as Penaeopsis challengeri belongs to P. eduardoi, as do the male from sta 12, the female from sta 38, and the 3 females from sta 74 which he named Parapenaeus rectacutus. Contrary to de Man’s statement, the juvenile does possess exopods on all pereopods; however, they are minute and thus easily overlooked. The descriptions and illustrations presented by Kubo (1949) leave no doubt that the Japanese shrimps he recognized as Penaeopsis rectactus (incorrect subsequent spelling of rectacutus which, according to Article 33b of the International Code of Zoological Nomenclature, cannot be used as a replacement name) are P. eduardoi. Kubo’s observations indicate that his specimens exhibit slight differences from those recorded by other authors, e.g., the sixth abdominal somite lacks a cicatrix whereas in those from other waters it possesses an interrupted one, and the length of the third antennular article is 1.5 times that of the second whereas in the material examined by me it is only about 1.2 times. Regarding his conclusion that the Japanese specimens differ from Indonesian specimens in possessing a postrostral carina, I wish to point out that it is based on an omission in the otherwise fine descriptive account given by de Man; such a carina is present in the specimens de Man saw. I have found the following slight intraspecific variation in thelycal fea- tures of P. eduardoi: The plate of sternite XIV may have the anterior and lateral borders joined forming an almost perfect arc or an obtuse angle, and the median plate of sternite XIII may vary from nearly semicircular to subcordiform. I have observed too that the third maxilliped, the third and, sometimes, fifth pereopods reach about the same level, but the fifth pereopod may not extend nearly so far anteriorly. Also, as previously indicated by Kubo (1949), the third maxilliped and the third pereopod are slightly longer in females than in males, and the ratio of dactyl:propodus of the third maxilliped is larger in the former than in the latter. These differences, however, are virtually insignificant. Acknowledgments I am deeply grateful to A. A. Fincham and R. W. Ingle, both of the BMNH, as well as to J. H. Stock of the ZMA for making available to me critical specimens under their care. H. H. Hobbs, Jr., made valuable sug- guestions to the manuscript, which also was improved by the critical read- ing of F. A. Chace, Jr., and A. B. Williams. Manuela Dieguez prepared the detailed illustrations. Literature Cited Alcock, A., and A. R. S. Anderson. 1894. An account of a recent collection of deep sea Crustacea from the Bay of Bengal and Laccadive Sea. Natural history notes T e . ‘ . ‘ ~ . } } from H. M. Indian Marine Survey steamer Investigator, Commander C. F. Old 182 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON ham, R. N., commanding. Series II, No. 14. Jour. Asiatic. Soc. Bengal 63, pt. 2: 141-185, pl. 9. Balss, H. 1925. Macrura der Deutschen Tiefsee-Expedition, 2. Natantia, A. Wiss. Ergebn. Deutsch. Tiefsee Exped. Valdivia 20:217-315, fig. 1-75, pl. 20-28. Bate, C. S. 1881. On the Penaeidea. Ann. Mag. Natur. Hist., ser. 5, 8:169-196, pl. 11-12. ——. 1888. Report on the Crustacea Macrura collected by H.M.S. Challenger during the years 1873-76. Rep. Sci. Res. Voyage Challenger, Zool. 24, xc + 942 DO) fel DO! pl: Burkenroad, M. D. 1934. Littoral Penaeidea chiefly from the Bingham Oceanographic Collection. With a revision of Penaeopsis and descriptions of two new genera and eleven new American species. Bull. Bingham Oceanogr. Collect. 4(7):1- 109, fig. 1-40. Ivanov, B. G., and A. M. Hassan. 1976. Penaeid shrimps (Decapoda, 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. Crustaceana 31:1—10, fig. 1-3. Kubo, I. 1949. Studies on penaeids of Japanese and its adjacent waters. Jour. Tokyo Coll. Fish. 36(1):1—467, fig. 1-160. Man, J. G. de. 1911. The Decapoda of the Siboga Expedition, Part 1. Family Penaeidae. Siboga Exped. Monogr. 39a, 131 p. —. 1913. The Decapoda of the Siboga Expedition, Part 1. Family Penaeidae. Siboga Exped. Monogr. 39a, (Suppl.), 10 pl. Pérez Farfante, I. 1969. Western Atlantic shrimps of the genus Penaeus. U.S. Fish Wildl. Serv., Fish. Bull. 67:461-591, fig. 1-77. Villaluz, D. K., and F. J. Arriola. 1938. Five other known species of Penaeus in The Philippines. Philippine Jour. Sci. 66:35-41, pl. 1-4. Yokoya, Y. 1933. On the distribution of decapod crustaceans inhabiting the con- tinental shelf around Japan, chiefly based upon the materials collected by S. S. Séy6-Maru, during the year 1923-1930. Jour. Coll. Agr., Tokyo Imp. Univ. 12: 1-296, fig. 1-71. —. 1941. On the classification of penaeid shrimps by the structural features of the appendix masculina. Jour. Coll. Agr., Tokyo Imp. Univ. 15:45-68, fig. 1-8, pl. 1-2. National Marine Fisheries Service, Systematics Laboratory, National Museum of Natural History, Washington, D.C. 20560. PROC. BIOL. SOC. WASH. 90(1), pp. 183-188 NEOPOMACENTRUS MIRYAE, A NEW SPECIES OF POMACENTRID FISH FROM THE RED SEA M. Dor and Gerald R. Allen It has come to our attention that one of the commonest pomacentrids in the Gulf of Aqaba, northern Red Sea is undescribed. The species is well known and usually referred to as Abudefduf azysron (Bleeker). However, in a monographic review of the Pomacentridae currently in progress, the junior author finds that azysron (now included in Neopomacentrus ) is con- fined primarily to the Malaysian-Indonesian Archipelago and adjacent areas. In most recent classifications, Neopomacentrus miryae would be placed in the genus Abudefduf on the basis of the smooth preopercle margin and single row of teeth in the upper and lower jaws. Allen (1975) pointed out that the previous classification of the Pomacentridae has been grossly over- simplified, particularly concerning Abudefduf and Pomacentrus. These two groups are separable into at least 13 genera, including Neopomacentrus Allen (1975). The latter genus is characterized by the following combi- nation of features: body relatively elongate, depth usually 2.3-2.8 in standard length; scales large, about 27-29 in a median lateral series; edge of preopercle smooth to moderately serrate; margin of suborbital usually smooth to weakly serrate or hidden by scales; teeth at front of jaws biserial in most species; dorsal rays XIII,10-13; middle rays of soft dorsal and anal fins, and outer rays of caudal fin frequently produced into elongate filaments. Measurements were made with dial calipers to the nearest one-tenth millimeter. Standard length is abbreviated as SL. Counts and proportions appearing in parentheses in the description section apply to the paratypes if differing from the holotype. Type-specimens have been deposited at the following institutions: Bernice P. Bishop Museum, Honolulu (BPBM); British Museum (Natural History ), London (BMNH); Hebrew University, Jerusalem (HUJ); Museum Na- tional d'Histoire Naturelle, Paris (MNHN); Tel Aviv University, Israel (TAU); National Museum of Natural History, Washington (USNM); Western Australian Museum, Perth (WAM). Neopomacentrus miryae, new species Rises Tabless 2 Holotype —BPBM 20322, 76.1 mm SL, collected with spear and quin- aldine in 10 m at Dahab, Sinai Peninsula, Gulf of Aqaba, Red Sea by J. Randall, 22 September 1974. 184 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Paratypes—BPBM 14327, 75.5 mm SL, collected at Elat, Gulf of Aqaba, Red Sea by H. Fricke, December 1972; BPBM 14666, 79.0 mm, same data as BPBM 14327 except collected 8 November 1972; BPBM 18208 (2 speci- mens, 63.4 and 79.3 mm SL), collected with the holotype; BMNH 1976.- 5.4.1-2 (2, 42.3 and 56.7), collected with spear in 15 m at Elat, Red Sea by G. Allen, 24 November 1975; HUJ 4825 (3, 38.0-86.0), collected with rotenone near Elat, Red Sea by E. Clarke, 22 July 1968; MNHN 1976-8, 64.5 mm, collected at Elat, Red Sea by H. Fricke, 8 November 1972; TAU 3225 (11, 58.0-83.6), collected at Elat, Red Sea with net by L. Fishelson, January, 1969; USNM 215463 (2, 82.2 and 86.0), collected at Elat, Red Sea, 2 October 1965 (collector unknown); USNM 216432 (31, 21.0-76.0), collected with rotenone in 3-12 m just north of Ras Burga, Gulf of Aqaba, Red Sea by V. Springer, 23 July 1969; WAM P25523-002, 45.0 mm, collected in 15 m at Elat, Red Sea by G. Allen, 24 November 1975. Diagnosis.—A species of Neopomacentrus with the following combination of characters: dorsal rays XIJJ,11-13; anal rays II,11; pectoral rays 19; tubed lateral line scales 17; gill rakers on first arch 27-30; edge of suborbital hidden by scales; caudal fin pale without dark markings; in life a prominent white spot just behind posteriormost dorsal rays. Description.—Dorsal rays XIII,12 ( XIII,11-13); anal rays II,11; pectoral rays 19 (18-19); branched caudal rays 13; gill rakers on first branchial arch 8 + 20 = 28 (7-9 + 19-21); branchiostegal rays 6; lateral line scales with tubes 17 (17-19); vertical scale rows from upper edge of gill opening to caudal base 29 (28-29); scales above lateral line to middle of spinous dorsal fin 1%; scales below lateral line to origin of anal fin 9 (9-10); circumpeduncular scales 16 (15-16). Body relatively elongate, depth 2.6 (2.2-2.6) in standard length, com- pressed, width 2.3 (2.3-2.7) in depth; head length 3.6 (3.2-3.7) in standard length; snout 4.5 (4.0-4.5) in head; eye 3.4 (3.2-3.8) in head; width of bony orbit 3.3 (2.9-3.5) in head; interorbital space moderately convex, bony width slightly less than eye diameter; least depth of caudal peduncle 2.2 (2.1-2.5) in head; length of caudal peduncle 1.9 (2.0-2.4) in head. Mouth oblique, terminal, maxillary ending slightly posterior to a vertical through anterior edge of eye; teeth of jaws incisiform, uniserial; upper jaw with 42 (38-42) teeth, largest about equal to diameter of nostril in height; lower jaw with 36 (34-36) teeth, largest slightly smaller than upper teeth; single nasal opening on each side of snout; nostril with a very low fleshy rim; margin of preorbital entire; suborbital margin mostly hidden by scales, free only anteriorly; margin of preopercle entire; opercle margins entire except a single flattened spine on upper portion. Scales finely ctenoid; preorbital, snout tip, lips, chin, and isthmus naked; remainder of head and body scaled; suborbital with a single row of scales; three parallel rows of scales below this to lower margin of pre- 185 VOLUME 90, NUMBER 1 ‘(qjepury oy of Aq oyoyd) vag poy “qeyeq “TS uur PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 186 ‘poseureqd , GGV LLE 90P O8E L6G 61€ uly [epneo jo Y}suoT 00G CST GIG OIG LZ TILT Aei [euB }sosUuo], JO YSU] 9ST GGT 8ST LOT LET GST ourds yeue pug jo yIsu0T 9) 192 1! 12, 69 99 oulds yeue 4ST Jo suo] iaal I&T CCT ial 8éI GSI oulds [esiop jsesuo] Jo ysus'T #L9 G6 C6 68 16 #09 oulds [esiop pug Jo yysueT vs 9¢ Me IV (as) 89 oulds [esiop js—T jo yysueT SLI CFI imal SPI SZI Ist euids otajed jo y}sue'T 68Z CVG 89% 69G 1G OIG Ulf OIAjod jo Wsus'T 60€¢ 00€ POS 166 COG €9G Ulf [e1oyood jo Yysus'T I1G 636 OFS i 6 1G 61Z eseq uly [eue jo YWsu0T ggg G09 F8C G8¢ 8GG 9LG oseq Ulf [eS1Op JO Y}suUNT DEP OLE G6E SIP O9€ CLE Ulf OLATed JO UISIIO 0} yNoOUS 9F9 669 LG9 8G9 Gr9 19 uly [PUB JO UISIIO 0} JnoUsg 9c¢ eGe 19€ LOE CSE cee Ulf [WSLOp JO UISIIO 0} JnoUS I€T CET PFI 6ET PET CFI gjounped [epneos jo ysueTy LET 9€T O€T inal 861 96T gpunped jepnes jo yjydep ysvoT 08 SL v8 06 CL GL [eHqtoroyur Auoq JO YIPIAA LOT 66 £6 98 18 G8 WqIo Auoq Jo JajyourKIG 69 69 TL GL c9 €9 Ysus] nous ITE 866 C66 662 CLE 18Z yyBuey prop] O9T 6ST GLI £06 COT ILI Apoq JO YPIM ysayea1y I6€ OCF OIP 9GP 88E c8e Apoq jo ydep ysayee1g (SP L9G cS 2°) 0'6L 0°98 T'9L (UIUL) YJsue, prepuRisg od A}e1e od A}eleg od Ayeieg od A}eleg od A}eledg ad A}0[0F{ Japeleyy GO0-ESSSSd CTVG9L6T S-9L6TNHNW 999FT Nd S9PSIGIWNSN GdE0s Wadd NVM HNW4d nnn nnn aT eee ‘(Yjsusey plepuejs Jo sy}pursnoy} ut) aphimu sniquaonuodoan jo suowitoods-od4} Jo suonsodoid o1njowoydioyy *T qe L VOLUME 90, NUMBER 1 187 opercle; dorsal and anal fins with a basal scaly sheath; caudal fin scaled nearly % distance to end of lobes; paired fins scaled only basally; axillary scale of pelvic fins about % length of pelvic spine. Tubes of lateral line ending below anterior rays of soft portion of dorsal fin; 4 (1-4) pored scales posterior to tubed scales; a series of 9 (7-9) pored scales mid-laterally on caudal peduncle to caudal base. Origin of dorsal fin at level of fourth tubed scale of lateral line; spines of dorsal fin gradually increasing in length to last spine (or middle spines in specimens under about 50 mm SL); membrane between anterior spines moderately incised, that between posterior spines only slightly incised; longest dorsal spine 2.3 (2.0-2.2) in head; first dorsal spine 1.2-1.6 in second spine (damaged in holotype); second dorsal spine 1.4-1.6 in longest dorsal spine; longest (6th) soft dorsal ray 1.5 (1.2-1.6) in head; length of base of dorsal fin 1.7 (1.7-1.8) in standard length; first anal spine about equal to first dorsal spine, its length 2.4 (2.2-3.0) in second spine; second anal spine 1.8 (1.8-2.0) in head; longest (7th) soft anal ray 1.6 (1.2-1.6) in head; base of anal fin 2.6 (2.4-2.6) in base of dorsal fin; caudal fin forked, its length 3.1 (2.6-3.4) in standard length; pectoral fin relatively short, not reaching a vertical through origin of anal fin in adults, longest ray 3.5 (3.2-3.8) in standard length; pelvic fin of adult usually not reaching origin of anal fin, longest ray 4.8 (3.5-4.5) in standard length. Color in 70% ethanol.—Holotype head and body brown, grading to tan ventrally; dark brown spot smaller than pupil size at uppermost portion of pectoral base, scarcely invading axil; median fins tan to slightly dusky; narrow black margin on dorsal fin; paired fins whitish. Paratypes similar in color except dark spot on base of right pectoral fin of largest USNM specimen expanded, forming a narrow bar on fin base. Color in life —Based on a 35 mm transparency taken by the junior author in 15 m at Elat. Head and body mostly olive green with dusky patch on each scale giving overall reticulated appearance; snout suffused with yellow; upper half of caudal peduncle and outer margins of lobes of caudal fin yellow-orange; prominent white spot, smaller than pupil, just behind posteriormost dorsal ray; dorsal fin grayish-green with narrow black margin; anal and pelvic fins whitish; pectoral fins transparent with small black spot on upper edge of fin base. Remarks.—This species appears to have no close relatives. It is the only species of Neopomacentrus lacking an inner row of slender buttress teeth at the front of the jaws. N. miryae is extremely abundant in the Gulf of Aqaba. Perhaps it is the most common pomacentrid found there. The ecology of this species was discussed by Fishelson et al. (1974) who incorrectly referred to it as Abudefduf azysron (Bleeker). Although azysron is a distinct species of Neopomacentrus, it does not occur in the Red Sea; its main distribution is 188 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Table 2. Frequency distribution for selected counts of types of Neopomacentrus miryde. Soft dorsal rays Pectoral rays 1i 12 13 18 19 3 All il 8 16 Gill rakers Lateral-line tubes upper lower ihe 18 19 1 8 9 19 20 Dil 14 9 If 1 18 D 6 3) 5 in the Malaysian-Indonesian Archipelago and adjacent regions (Allen, 1975). Baschieri-Salvadori (1957) was the first to use this name for a Red Sea pomacentrid, however, the fish which he called azysron was not N. miryae, but a second member of the genus, inhabiting the southern Red Sea. It will be described in a review of Red Sea pomacentrids cur- rently in preparation by the junior author and J. E. Randall. We have named the new species miryae in honor of the senior author's late wife. Acknowledgments We would like to express our gratitude to Mr. David Fridman, Manager of Coral World, and Dr. Ilan Paperna, Director of the Hebrew University Marine Laboratory for providing facilities during the junior author's visit to Elat during November 1975. Thanks are also due to Dr. J. E. Randall for sending specimens and a photograph of the type of N. miryae. Literature Cited Allen, G. R. 1975. Damselfishes of the South Seas. T. F. H. Publications, Inc., Neptune, New Jersey. 240 pp. Baschieri-Salvadori, F. 1957. Spedizione subacqua Italiani nel Mar Rosso Ricerche Zoologiche. Ix. Pomacentridi. Riv. Biol. Colon. 15:57-68. Fishelson, L., D. Popper, and A. Avidor. 1974. Biosociology and ecology of poma- centrid fishes around the Sinai Peninsula (northern Red Sea). Jour. Fish. Biol. 6:119-133. College of Kibbutz Education, Tel Aviv, Israel, and Western Australian Museum, Perth, Australia. 71 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, Washing- ton, D.C. 20560. Review.—One of the Society’s aims is to give its members an opportunity for prompt publication 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 international 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 814 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 indica- tions 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 Faxonella blairi, a new crawfish from the Red River drainage of Oklahoma and and Arkansas. William A. Hayes and Rollin D. Reimer Percina (Imostoma) antesella, a new percid fish from the Coosa River system in Tennessee and Georgia. James D. Williams and David A. Etnier Apogon mosavi, a new western Atlantic cardinalfish, and a note on the occurrence of Apogon leptocaulus in the Bahamas. George Dale Description of Colomastix janiceae n. sp., a commensal amphipod (Gammaridea: ° Colomastigidae) from the Florida keys, USA. Richard W. Heard and Dan Perlmutter Sarsiella pseudospinosa, a new marine ostracod (Myodocopina; Sarsiellidae) from southern California. James H. Baker Description of a new subspecies of prairie vole, Microtus ochrogaster. William D. Severinghaus Ophiacantha clypeata n. sp. from the Bering Sea, with a redescription of Ophiacantha rhachophora Clark (Echinodermata: Ophiuroidea). Michael A. Kyte A new species of Eleutherodactylus (Anura: Leptodactylidae) from the Cordillera Oriental of Colombia. William E. Duellman and John E. Simmons Spionidae (Annelida: Polychaeta) from San Francisco Bay, California: A revised list with nomenclatural changes, new records, and comments on related species from the northeastern Pacific Ocean. William J. Light A new genus of Pacific Etelinae (Pisces: Lutjanidae) with redescription of the type-species. William D. Anderson, Harry T. Kami, and G. David Johnson A synoptic list of the described ground beetle larvae of North America (Coleoptera: Carabidae). Raymond G. Thompson Cave shrimps in the Caicos Islands. Donald W. Buden and Darryl L. Felder An emendation of the genus Sathrodrilus Holt 1968 (Annelida: Branchiobdellida), with the description of four new species from the Pacific drainage of North America. Perry C. Holt Biological results of the University of Miami deep-sea expeditions. 121. A review of the recent species of Balanophylla (Anthozoa: Scleractinia) in the western Atlantic, with descriptions of four new species. Stephen D. Cairns A new species of pygmy characoid fish from the Rio Negro and Rio Amazonas, South America (Teleostei: Characidae). Stanley H. Weitzman and Robert H. Kanazawa A new marine genus and species of the Nuuanu-group (Crustacea, Amphipoda) from the Yucatan Peninsula. Larry D. McKinney and J. Laurens Barnard Penaeopsis eduardoi, a new species of shrimp (Crustacea: Penaeidae) from the _ Indo-West Pacific. Isabel Pérez Farfante Neopomacentrus miryae, a new species of pomacentrid fish from the Red Sea. M. Dor and Gerald R. Allen 19 30 43 49 55 60 66 89 og 108 116 132 > / woe 72 Proceedings of the BIOLOGICAL SOCIETY of WASHINGTON Volume 90 August 12, 1977 Number 2 THE BIOLOGICAL SOCIETY OF WASHINGTON 1977-1978 Officers President: Richard S. Cowan Secretary: W. Duane Hope Vice President: Clyde F. E. Roper Treasurer: Oliver S. Flint, Jr. Custodian of Publications: John H. Miles Elected Council J. Laurens Barnard Raymond B. Manning Ronald I. Crombie John H. Miles Frank D. Ferrari PROCEEDINGS Editor: Austin B. Williams Coeditor: C. W. Hart, Jr. f Associate Editors Classical Languages: George C. Steyskal Invertebrates: Thomas E. Bowman Plants: David B. Lellinger Vertebrates: Leslie W. Knapp 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 Institution, Washington, D.C. 20560. The Proceedings of the Biological Society of Washington is issued quarterly. Manuscripts, corrected proofs, editorial questions should be sent to the Editor, Biologi- cal Society of Washington, National Museum of Natural History, Smithsonian Institution, Washington, 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 Application to mail at second class postage rates is pending at Washington, D.C., and — additional mailing office. : PROG? BIO, SOG. WASEL- 90(2), pp. 189-193 REDESCRIPTION OF SALMINCOLA LONGIMANUS GUNDRIZER, 1974 (COPEPODA: LERNAEOPODIDAE) Z. Kabata In his study of parasites of fishes occurring in the waters of the Tuva ASSR, in Siberia, Gundrizer (1974) described a new species of Salmincola Wilson, 1915. This new species, S. longimanus, was found in the nostrils of the Siberian grayling, Thymallus brevirostris Kessler, taken in Lake Mumuday situated in the drainage area of the river Kobdo. The main distinguishing characteristic of S. longimanus is the length of its second maxillae, a feature which prompted the specific name of the parasite. In addition to the habitus drawing, Gundrizer’s description included only a generalized drawing of the maxillipeds. No description of appendages was given in the text. My Russian colleagues asked me to examine and redescribe Gundrizer’s material and sent me 2 specimens. Both were female, one ovigerous. With the aid of the same methods as those used by me earlier in the revision of the genus Salmincola (cf. Kabata, 1969), I examined these 2 specimens in some detail. The results of my examination are described and illustrated below. Salmincola longimanus Gundrizer, 1974 Figs. 1-12 Female.—Cephalothorax inclined ventrally to long axis of trunk (Fig. 1), long and tapering (maxillipeds not extending to level of cephalothorax tip) (Fig. 2). Definite, though shallow, constriction behind bases of second maxillae. Trunk narrower anteriorly, expanding posteriorly into almost regularly hemispherical extremity. In preserved specimens trunk somewhat flexed ventrally. Egg sacs relatively short and bulky. Dimensions of speci- mens examined (in mm) as follows: Cephalothorax length Oo 2.07 Trunk length 3.40 232 Trunk width (anterior ) 1.00 0.66 Trunk width (greatest) Sa 1.49 Second maxillae, length 4.98 4.65 Second maxillae, width 0.50 0.41 Egg sacs, length 2.24 — Egg sacs, width 1.08 — First antenna (Fig. 3) digitiform, obscurely segmented, with apical armature reduced or absent (probably due to damage in vivo). Second antenna (Fig. 4) with broad, 2-sezmented sympod, 2-segmented endopod 190 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Vy vy VVyyt y v ny) Voy Figs. 1-7. Salmincola longimanus, female. 1, Entire, lateral view; 2, Entire, dorsal view; 3, First antenna, ventral; 4, Second antenna, entire, lateral; 5, Same, endopod, lateral; 6, Same, different specimen, medial; 7, Tips of second maxillae and bulla. (Explanations in text. ) and l-segmented exopod; second segment of sympod with raised lateral pad bearing batch of slender spinules; proximal segment of endopod fused with sympod, on its ventral wall round, inflated and spinulated pad [spinules of pad either slender (Fig. 5) or minute (Fig. 6) and more nu- merous]; distal segment with usual apical armature; in smaller specimen VOLUME 90, NUMBER 2 190 eecO ain, Figs. 8-12. Salmincola longimanus, female. 8, Mandible, lateral; 9, First maxilla, lateral; 10, Maxilliped, entire, dorsal; 11, Same, palp, ventral; 12, Same, tip of sub- chela, dorsal. (p—palp. ) process 4 unarmed and prominent, longer than hook 1 (Fig. 5), in larger specimen process 4 smaller and armed with tiny spinules (Fig. 6), both with auxiliary spine 2 and small ventral process 5 (not seen in Fig. 6), but tubercle 3 not observed; exopod indistinctly divided into soft lobes at apex, bearing either slender or very minute spinules and lateral papilliform process. Mandible (Fig. 8) with 7 teeth, proximal 3 distinctly smaller than others, overlapped by ventral blade (in one mandible pressure on coverslip pushed blade away and exposed 3 basal teeth). First maxilla (Fig. 9) with small, conical exopod surmounted by spiniform process; 3 endopod papillae short, carrying almost conical terminal setae, ventral papilla noticeably smaller than other 2; appendage tapering distally, its greatest width proximal to level of exopod. Second maxilla cylindrical (Fig. 1), 192 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON of uniform diameter except at expanded tips (Fig. 7). Bulla (Fig. 7) regularly obovate, with short manubrium. Maxilliped (Fig. 10) broad at base, tapering distally; ill-defined transverse wrinkles possibly denoting obsolete segmental boundary; distal to this boundary and close to medial margin small, soft palp (Fig. 11) with apparently bifid tip; base of sub- chela unarmed, its apex with inflated double pad bearing numerous, blunt, minute denticles (Fig. 12); claw greatly reduced. Discussion The specimens described above are in fairly good agreement with Gundrizer’s (1974) general description. Their dimensions also differ only a little from the size range given by the discoverer of the species. I was, however, unable to see a short, tapering outgrowth on the medial margin of the maxilliped proximal to the palp. Such outgrowth was shown by Gundrizer in his Fig. 2. I know of no such outgrowth in the maxillipeds of any species of Salmincola and cannot account for it. It was interesting to find that the 2 specimens examined differed from each other in the armature of the second antennae (Figs. 5 and 6). This suggests that the species has a fairly broad range of morphological vari- ability. It is even more interesting, when one recalls that Salmincola thymalli (Kessler, 1868), parasitic exclusively on Thymallus, is also dis- tinguished by similar variability (Kabata, 1969). Differences in the armature of the second antenna, occurring within the same population, were also recorded by Kabata and Koryakov (1974) in Salmincola cottidarum Messjatzeff, 1926, in Lake Baikal. Gundrizer (1974) described also a subspecies of S. longimanus, which he designated S. longimanus sibiricum and which differed from the typical form in having a spherical bulla and strange outgrowths on the maxilliped, at the base of the subchela. Should the presence of these structures be corroborated by later investigations, they too, will be unique to the genus and, indeed, to Lernaeopodidae. Keeping in mind the morphological variability of S. longimanus, as represented by differences between the 2 specimens examined, I doubt whether it is necessary to accord them a sub- specific rank. S. longimanus is clearly an interesting species and a thorough study of its morphology and biology might contribute significant new details to our knowledge of Lernaeopodidae. S. longimanus, by virtue of the reduced claw of its maxilliped, belongs to the subgenus Salmincola (Brevibrachia) Kabata, 1969. The members of this subgenus are mainly parasitic on Coregonidae, so that S. longimanus is rather less than a typical member. Its addition to the subgenus calls for some modifications to the key first published by Kabata (1969), and constituting a part of the key to the entire genus. VOLUME 90, NUMBER 2 193 1. Bulla with moderately long manubrium and flat subcircular anchor very wide in diameter; process 4 on endopod of second antenna always much smaller than hook 1, unarmed; egg-sacs twisted in anterodorsal direction S. extumescens Bulla with short manubrium and anchor other than flat and sub- circular 2, 2. Bulla irregular, its anchor hoof-shaped S. strigatus Bulla obovate, second maxillae as long as or much longer than trunk, subchela of maxilliped inflated at apex, bearing batch of blunt, minute denticules S. longimanus Bulla more or less spherical, with short manubrium 3 3. Second maxillae shorter than trunk; subchela of maxilliped without denticles at base of claw; trunk with 3 swellings on lateral margins S. nordmanni Second maxillae shorter than trunk; subchela of maxilliped with denticles at base of claw; lateral margin of trunk smooth S. extensus and S. jacuticus* Acknowledgment I wish to express my gratitude to Dr. A. V. Gusev and Dr. O. N. Bauer, both of Leningrad, USSR, who kindly supplied me with the material used in this paper. Literature Cited Gundrizer, A. N. 1974. Paraziticheskie veslonogie ryb Tuvy. [Parasitic copepods of fishes of Tuva.] Trudy nauch.-issled. Inst. Biol. Biofiz. Tomsk. gos. Univ., Biologiya 3:61-68. Kabata, Z. 1969. Revision of the genus Salmincola Wilson, 1915 (Copepoda: Lemaeo- podidae). Jour. Fish. Res. Board Canada 26:2987-3041. , and EK. A. Koryakov. 1974. Morfologicheskaya izmenchivost Salmincola cottidarum Messjatzeff, 1926 (Copepoda: Lernaeopodidae), parazita bychkov. [Morphological variability of Salmincola cottidarum Messjatzeff, 1926 ( Cope- poda; Lernaeopodidae), a parasite of Cottocomephorinae.] Parazitologiya 8: 306-311. Department of the Environment, Fisheries and Marine Service, Pacific Biological Station, Box 100, Nanaimo, B.C., Canada V9R 5k6. *For reasons why these two species cannot be distinguished from each other see Kabata (1969). PROC. BIOL. SOC. WASH. 90(2), pp. 194-204 A NEW SPECIES OF FROG OF THE GENUS ELEUTHERO- DACTYLUS (AMPHIBIA: LEPTODACTYLIDAE) FROM HE COCKE COUNTRY SOE I ANIAIG: Ronald I. Crombie Among the more prominent geological features of Jamaica are the extensive jagged limestone hills of the Cockpit Country. The harsh terrain of this area has long discouraged herpetological collecting, but recently two new species have been described from the more accessible periphery of the Cockpits (Schwartz, 1971; Thomas, 1975). Since similar karst areas on other Greater Antillean islands harbor endemic faunas, these discoveries indicated that the Cockpits might also. With this in mind, Jeremy F. Jacobs and I walked several miles overland into the southwestern Cockpits in the summer of 1974. The trip in was arduous and not particularly pro- ductive, but while walking out we collected two specimens of a distinctive small Eleutherodactylus. We returned to the area for two days and a night in July 1975 and collected several more specimens. We then walked the remaining distance across the Cockpits, from south to north, sampling as many different habitats as possible before emerging near Deeside. Eleutherodactylus sisyphodemus, new species Fig. 1 Holotype.—National Museum of Natural History (USNM) 200000, a gravid female from the vicinity of “the cave” about 4 mi WNW Quick Step, Trelawny Parish, Jamaica, collected by Jeremy F. Jacobs and R. I. Crombie, 12 July 1975. Paratypes (4)—Museum of Comparative Zoology (MCZ) 89138, same data as holotype; USNM 200001, same locality as holotype, collected by Barbara A. Harvey and Jeremy F. Jacobs, 12 July 1975; Albert Schwartz Field Series (ASFS) V28447, same locality as holotype, collected by R. I. Crombie, 12 July 1975; USNM 200002, on trail between “the cave” and the road north of Quick Step, Trelawny Parish, Jamaica, collected by J. F. Jacobs and R. I. Crombie, 29 August 1974. Referred specimens (2)—USNM 200004, same data as holotype; USNM 200003, on trail between “the cave” and the road north of Quick Step, Trelawny Parish, Jamaica. Collected by J. F. Jacobs and R. I. Crombie, 29 August 1974 (cleared and stained). Diagnosis.——A small species (males to 13.8, females to 17.9 mm snout- vent length) that is easily distinguished from all other Jamaican Eleuthero- dactylus by the presence of a calcar and denticulate tarsal finge, limited dorsal pattern polymorphism, and the black ventral coloration. VOLUME 90, NUMBER 2 195 BY id > Fig. 1. Eleutherodactylus sisyphodemus, new species, holotype (USNM 200000): snout—vent length 17.9 mm. Description of holotype—A gravid female with the following measure- ments (mm): snout—-vent length (SVL) 17.9; head length (HL) 7.1; head width (HW ) 7.0; tympanum diameter (TYM) 1.5; eye diameter (EYE) 2.2; naris-eye distance (NE) 2.0; femur length (FEM) 8.2; tibia length (TIB) 8.1; foot length (FT) 8.2. Head slightly longer than broad; HW SVL 39.1%. In lateral view, snout protruding; canthus rostralis rounded; loreal region slightly concave; lips not flared. Nostrils lateral, much closer to tip of snout than eye; diameter of eye slighter greater than distance from naris to eye; interorbital distance 2.7, greater than eye diameter. A single prom- inent tubercle in center of each eyelid with several more indistinct ones posteriorly and along edge of eyelid. Tympanum round, distinct, not sexually dimorphic in size. Tongue lanceolate but not nicked behind, tree 196 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON for more than half its length posteriorly. Choanae small and round; pre- vomerine dentigerous processes in two long, arched series extending from level of inner choanal border almost to palatal midline; separated by a distance equal to diameter of a choana. Fingers relatively short, unwebbed, with faint lateral fringe; III, IV, II, I in order of decreasing length. Digital pads flattened, longer than wide and distinctly pointed, with a circum- ferential groove. Several large, flat palmar tubercles; basal subarticular tubercle enlarged and flattened; a single additional subarticular tubercle present only on fingers III and IV; no supernumerary tubercles; a single pale ulnar tubercle on each arm. Toes long, unwebbed; IV, III, V, II, I in order of decreasing length; distal subarticular tubercles flattened and enlarged except conical and small on toes IV and V; several low conical plantar tubercles; a pale, swollen inner metatarsal tubercle. Digital pads similar to those on fingers but slightly larger. Heels not quite touching when femora at right angles to body axis; TIB/SVL 45.2%. A prominent, fleshy triangular spur at metatarsal-tarsal articulation and a larger one at tibio-tarsal articulation, joined by a scalloped dermal fringe; a less distinct fringe on both anterior and posterior faces of tibia. Anus directly pos- teriorly, with a series of four pale warts and an irregular black blotch just above it. Skin of dorsum smooth with two vague scapular warts and several irregularly scattered smaller pustules especially on upper surfaces of hindlimbs. A glandular lateral ridge continues from eye to about midway between limb insertions where it breaks up into a series of low tubercles. Venter smooth to finely shagreened, becoming slightly areolate near vent and bases of thighs. Coloration of holotype.—tIn life the dorsal ground color was slate gray, slightly paler laterally, with a faint silver middorsal hairline. A wider golden stripe extended from tip of snout, through the nostril, across edge of upper eyelid, and onto the lateral ridge where it faded before reaching the groin. Below this stripe, the entire lateral surface of the head was black; this mask extended to and covered the tympanum, beyond which it narrowed to a sharp point near where the lateral ridge ended. The two scapular warts and the supra-anal tubercles were dirty white; there was a black smudge just above the row of supra-anal warts. Dorsal surfaces of the limbs were lighter gray, suffused with tan; thighs were banded with darker gray with irregular black patches at the knee. The forearm had a dorsal median black spot which extended onto the inner surface. A pale dot was present at the base of each digital pad, on both dorsal and ventral faces of digit. The venter was charcoal gray to black with irregularly shaped silver flecks; the larger flecks were concentrated in the pectoral area and on the upper arms and thighs. Undersides of the limbs were slightly lighter gray; palms of the hands were black with a silver spot at the base of each VOLUME 90, NUMBER 2 197 digit and undersides of the pads were gray. A wide black stripe covered the ventral surface of the lower arm, blending with the dorso-lateral spot. Posterior faces of the thighs were black, forming a triangular “seat patch” with its apex at the anus. The posterior edge of the tarses and soles of the feet were also black; ventral sides of the pads and tarsal fringe were contrasting lighter gray. There were no “flash colors” but the inguinal areas were lighter gray. The iris was coppery red. Morphological variation—Four gravid females (including the holo- type) have the following measurements (mm) and proportions: SVL 15.6-17.9 (x = 16.9); HL 6.5-7.1 (6.8); HW 6.0-7.0 (6.6); TYM 1.3-1.6 (lta) EVEN 2:0=2.2 (2:0); NE 1.8-2.1 (2:0); FEM 7.2-8.2 (7.6); TIB 7:3— 8.1 (7.6); FT 6.9-9.2 (7.5); TIB/SVL 44.0-46.8% (45.0); HW/SVL 38.5- 40.0% (39.1). The two adult males have the following measurements and proportions: SVL 12.0-13.8; HL 5.4-5.5; HW 4.8-5.6; TYM 1.3-1.7; EYE 1.6-1.8; NE 1.5-1.8; FEM 5.8-6.7; TIB 6.0-6.7; FT 5.5-6.4; TIB/SVL 48.6-50.0%; HW/ SVL 40.0-40.6%. Both have vocal slits. A single juvenile (USNM 200004) measures 7.9 mm SVL. Morphological differences in the sample are slight. The angle of the prevomerine teeth series ranges from very slightly to distinctly arched; subarticular tubercles on hands and feet are variably flattened or conical, possibly due to differences in mode of preservation. The character showing greatest variation is texture of the dorsal skin. Four specimens (MCZ 89138, ASFS V28447, USNM 200003, females; and USNM 200001, male) have prominent conical lumbar tubercles. Of these individuals, MCZ 89138 and USNM 200003 have additional distinct tu- bercles only at the scapula. USNM 200001 has paired, unpigmented scapu- lar and sacral tubercles plus scattered obscure warts on the lateral areas. ASFS V28447 has extreme development in rugosity; both lumbar and scapular warts are well developed and they are joined by two almost com- plete rows of para-vertebral tubercles. Other individuals (USNM 200000, 200002, and 200004) lack lumbar warts and have, at best, only vague additional dorsal ornamentation. However, the tarsal fringe is very distinct in all specimens, even the juvenile. Myology and osteology.—In light of recent hypotheses on the polyphy- letic nature of the genus Eleutherodactylus, one specimen (USNM 200003 ) of E. sisyphodemus was dissected and later cleared and stained to determine the state of characters used by Heyer (1975:33) in his analysis. State terminology of Heyer is given in parentheses. Jaw musculature: The depressor mandibulae consists of three distinct slips with origins on the fascia, squamosal, and annulus tympanicus (DFSQat, in the terminology of Starrett, 1968; State A of Hever) with the annulus tympanicus slip being distinctly reduced. In the adductor 198 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON musculature of the squamosal, only the a.m. posterior sub-externus is present (State B). Hyoid musculature: The geniohyoideus medialis is basically contiguous medially (State A), but with some very slight medial separation. The anterior petrohyoideus attaches on the lateral edge of the hyoid plate (State A). The sternohyoideus insertion is entirely near the lateral edge of the hyoid body (State A), but its attachment is very short and restricted to the region of the alary process. The omohyoideus inserts laterally on the posterolateral process of the hyoid (State I). Thigh musculature: The iliacus externus is long, extending almost to the tip of the ilium. The tensor fasciae latae inserts on the anterior end of the ilium immediately anterior to the iliacus externus and the two muscles are contiguous for a considerable length (State F). Interior and exterior parts of the semitendinosus are approximately equal, displaced and attached by a tendon (State D). The adductor longus is absent (State D). Cranial osteology: The quadratojugal is present and contacts the maxilla (State A); there is very slight contact between the nasals and fronto- parietal (State A). The otic ramus of the squamosal is much longer than the zygomatic ramus, and approximately as long as the ventral ramus (State G). There is extensive median contact of the prevomers (State C) but no prootic/frontoparietal fusion (State A). The only exceptional element in the skull is the broad, rather heavily ossified lateral expansion of the otoccipital, creating a surprisingly solid posterior cranium for such a small frog. Postcranial osteology: The posterior sternum has a partially mineralized mesosternum and a cartilaginous xiphisternum (State C; most similar to fig. 4-J in Heyer, 1975:18). The sacral diapophyses are rounded and the transverse processes of the presacral vertebrae are long and slender. The terminal phalanges are distinctly T-shaped and there are prominent humeral and ilial crests. Pattern polymorphism.—Jamaican Eleutherodactylus are known to ex- hibit a bewildering variety of dorsal colors and patterns (Goin, 1954, 1960 and papers cited therein). E. sisyphodemus seems comparatively con- servative in its variation. Since there are so few specimens involved, the pattern of each is discussed separately. Holotype, USNM 200000: This specimen exhibits the simplest pattern type, an essentially unicolor dorsum with faint middorsal hairline and supra-anal dark splotch. Previously discussed. USNM 200002, male: Similar to the holotype, with distinct hairline and small anal blotch but no other dorsal markings. USNM 200001, male: Distinct hairline, but lumbar warts are surrounded by an irregular dark spot (pelvic spot of Goin, 1960:256, fig. 7). There is no dark anal patch and the legs are indistinctly barred. VOLUME 90, NUMBER 2 199 USNM 200003, female: Similar to 200001 but with more distinct leg barring and a small anal patch. MCZ 89138, female: Faint hairline, distinct dark lumbar spots and paired, smaller supra-anal spots. There is very faint suggestion of a dorsal “picket” pattern (Goin, 1960:250, fig. 3), but the “picket” is slightly darker than surrounding ground color. Legs distinctly barred. ASFS V28447, female: Extremely faint hairline. A distinct dark “picket” outlined by tubercles. The lumbar spots are large and several outlining warts of the picket, including the scapular warts, are darkened. USNM 200004, juvenile: No hairline. A large, pale, sub-rectangular mid- dorsal patch. The lumbar spots are white. One other specimen (an adult that was lost on the cross-Cockpit hike) exhibited this pattern. Variation in color was slight. Dorsal ground color in life ranged from rich, milk chocolate brown (ASFS V28447) to the slate gray of the holotype. Ventral color was uniformly shiny dark gray to black. The number of silver flecks was variable, but most individuals had more extensive, more evenly distributed flecking than the holotype. The middorsal hairline (middorsal stripe of Goin, 1960:251, fig. 4) is common in West Indian Eleutherodactylus. Eleven of the 16 Jamaican species of the genus (8 of the 10 members of the gossei group) exhibit this morph (Goin, 1960; Schwartz and Fowler, 1973; Crombie, pers. obs.). It is a relatively consistent element of the pattern in all E. sisyphodemus, except the “dorsal blotch” morph. Of the three basic morphs in the sample, the picket pattern is relatively rare in Eleutherodactylus, it is found only in two members of the gossei- group (gossei and pantoni) and one mainland middle American species (Goin, 1960). The picket in sisyphodemus differs from the typical pattern in that the picket is darker than the surrounding ground color rather than lighter as in the other species exhibiting the morph. Goin (1960:256) reported without documentation that the pattern of bilaterally symmetrical “pelvic spots” (= lumbar spots) was “widespread in the genus.” In Jamaican species the morph is restricted to members of the gossei-group. E. andrewsi almost invariably has this pattern, but indi- viduals have been found with spots of differing size and shape, one spot, or no spots, although the latter is rare. E. gossei, nubicola, pantoni, and luteolus occasionally have some trace of lumbar spots and E. fuscus shows the morph more commonly (Schwarz and Fowler, 1973; Crombie, pers. obs. ). The pale “dorsal blotch” appears unique in Jamaican (and possibly all West Indian) Eleutherodactylus. Comparisons.—Eleutherodactylus sisyphodemus is the smallest Jamaican frog; this fact, coupled with the tarsal ornamentation will easily distinguish it from all other species. When the first specimens were collected, I noted their resemblance to E. andrewsi, a species restricted to moderate ele- 200 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON vations in the Blue Mountains, far to the east of the Cockpits. E. andrewsi frequently has a unicolored dorsum with dark lumbar spots and a heavily pigmented venter. However, E. andrewsi is a highly vocal frog, and the ventral ground color is usually light with an extensive dark reticulum or spotting. The slightly larger size, occasional pale reddish flash colors, absence of limb fringes, and isolated distribution further distinguish that species. E. sisyphodemus and E. luteolus are syntopic at the type-locality of the former. They are similarly sized, but adult luteolus usually have bright yellow bellies. Some luteolus juveniles are dark; one specimen (7.8 mm SVL) was originally mistaken for sisyphodemus. However, the lack of a black ulnar stripe and tarsal fringes distinguished it from the similarly sized (7.9 mm SVL) sisyphodemus juvenile; therefore, it is relatively easy to identify even recently hatched material. The only other species with which sisyphodemus could be confused is E. junori. The loud, distinctive voice of junori was not heard in the deep Cockpits, but I have collected the species in the southeastern Cockpits, northwest of Troy. E. junori is a small species (19-27 mm) but with bright red flash marks in the groin and the characteristically variable dorsal pattern of most gossei-group members (see Relationships). Relationships—I divide Jamaican Eleutherodactylus into the following four groups, arranged in approximate order of decreasing SVL (I = introduced species). gossei-group ricordi-group auriculatus-group jamaicensis-group pantoni cundalli johnstonei (1) jamaicensis nubicola cavernicola fuscus grabhami gossei planirostris (1) orcutti alticola andrewsi junori luteolus sisyphodemus Schwartz (1969) characterized the auriculatus-group as moderate-sized frogs with a granular belly, short patchlike vomerine series, and well developed digital pads; most are vocal and call from arboreal sites. Schwartz (1969:114) hesitantly listed E. jamaicensis as a Jamaican representative of the auriculatus assemblage but noted that it was inconsistent in many ways with his concept of the group. I concur with this evaluation, but feel that the differences warrant exclusion from the auriculatus-group. VOLUME 90, NUMBER 2 201 Schwartz (1958; 1969:102) attributed the following characters to the ricordi-group: rugose dorsum; long vomerine series; smooth or feebly rugose-venter; feeble digital pads or enlarged ones restricted to the two outer fingers; faint, insect-like call. The four species listed above fit well within this grouping; therefore, I do not recognize the “cundalli group” of Goin (1954:185). Goin (1954:185) defined the gossei-group as follows: feebly developed digital pads; smooth back and belly; long vomerine series; red flash colors in the groin and on concealed portions of legs; vocal frogs with terrestrial calling sites. The group is somewhat more heterogeneous than Goin im- plied, but with the addition of two species my arrangement agrees with his. Goin felt that absence of red flash colors in E. luteolus warranted its exclusion from the gossei-group, but luteolus occasionally has red or orange pigment in the groin and inguinal areas. In addition, Goin overlooked the fact that bright colors were variable in other species he included in the group (absent in nubicola, present or absent in gossei and andrewsi). I tentatively follow Schwartz and Fowler (1973:126) in considering E. orcutti a specialized member of the gossei assemblage. This species has webbed feet, large digital pads, rugose dorsum, and no flash colors, all of which were interpreted as adaptations to a streamside niche. E. sisyphodemus is similarly specialized for existence in a restricted habi- tat. The species resembles other gossei-group members with its squat body shape, short legs, and long vomerine series, but the tuberculate dorsum, moderately well developed digital pads, and tarsal fringe are rare or un- known in the group. Most of these characters are adaptive for the forest floor-leaf litter niche the species occupies. Pending further studies, I consider E. sisyphodemus a somewhat anomalous member of the gossei- group. Natural history—The steep ridges of the Cockpit Country are part of the “White Tertiary or Quaternary limestone formation” (Versey, 1972). These ridges are often greatly eroded from the effects of heavy rainfall (100-150 inches/year) and underground rivers; sinkholes and large soil- filled solution pockets are common and huge boulders or rock outcroppings litter the slopes. Footing is precarious since the limestone is brittle and breaks off in hand or underfoot. A thick layer of wet leaves further hinders walking and often conceals deep sinkholes or other natural pitfalls. The vegetation of the Cockpits is “wet limestone forest” (Asprey and Robbins, 1953). The slopes are covered by a heavy growth of saplings interspersed with occasional larger hardwood trees (Cedrela, Brosimum, Nectandra); the canopy is complete and about 20-40 feet in height. The undergrowth is not thick and is primarily composed of climbing aroids, ferns, lianas and other epiphytes. Tank bromeliads are not especially 202 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON common in the understory but are more numerous high in the canopy or on exposed outcroppings. The valleys or Cockpits between the ridges are normally thick with low vegetation and larger trees but most of the more accessible valleys have been cleared for cattle grazing and/or cultivation. In the distant Cockpits, care of crops is sporadic and heavy secondary growth of wild plantain, wild yam, thatch palm, and other feral plants is common. About two miles north of Quick Step on the as yet unfinished cross- Cockpit highway is a small road leading to a lean-to, beyond which is a small agricultural area. An obscure path leads through a corn field, a small grove of citrus, and then into the shaded coolness of the Cockpit forest. “The cave” is well known to residents of southern Trelawny, but it is infrequently visited because of local lore concerning the somewhat truculent spirits that inhabit the area. The cave is on the third ridge, about two miles by foot from the lean-to. The valley at the base of the cave ridge is badly overgrown and hasn't been grazed for several years; the ridge slopes gently at the bottom but becomes progressively steeper and more boulder- strewn as one nears the caves mouth. All of the specimens of E. sisyphodemus were taken in the heavy leaf litter on this or similar ridges. The first 1974 specimen was collected as it hopped across the trail on the second ridge; it apparently had been dis- turbed by the first people in the line. The second specimen was found on the opposite side of the same ridge in the litter of a bromeliad we had dismantled. We were unsure whether it had come from the bromeliad or had been aroused from the leaf litter by the activity. In 1975, all the speci- mens were taken on the cave slope. One female was found crossing the trail during the day as we first ascended towards the cave mouth. After dark we collected several more specimens (mostly lost later on the trek through the Cockpits) by shuffling through a pocket of leaf litter until a frog came out. No call was heard that could be attributed to E. sisyphodemus but the choruses of other species were so intense that a quiet call could easily have been overlooked. Eleutherodactylus pantoni was the dominant voice in the nocturnal din, but the soft call of E. luteolus was also common. An irregular, high-pitched whistling tentatively traced to E. grabhami was unlike that recorded for the species elsewhere in its range. E. gossei was absent from the forest and only a few individuals were calling from the valley. A cundalli-like Eleutherodactylus was found in seepage areas of the cave and one large female was collected just outside the cave mouth on a leaf. Other anurans participating in the nocturnal chorus were: Hyla wilderi, Osteopilus brunneus and Calyptahyla crucialis; several unfamiliar calls were also heard. VOLUME 90, NUMBER 2 203 No reproductive activity was noted in the field but all females collected contained large, unpigmented eggs; USNM 200003 contained six ova (0.5-1.0 mm diameter) in the left oviduct and three smaller ones in the right oviduct. The stomach contents of this individual contained the remains of ants and a winged hymenopteran. The cryptic, leaflike body shape of E. sisyphodemus is reminiscent of several species of forest-floor dwelling leptodactylid and microhylid frogs. The cryptic coloration, absence of flash markings, and the tarsal decoration are obviously additional adaptations for life in the leaf litter community. Since this distinctive species remained undiscovered for so long, the possi- bility exists that it may be ecologically restricted to well shaded, undis- turbed, wet limestone forest. This hypothesis was supported by considerable collecting in slightly disturbed parts of the Cockpits where leaf litter was present but subject to periodic drought due to the incomplete canopy of the disturbed forest. Intensive search in these areas yielded only E. luteolus, pantoni, and grabhami. I anticipate that E. sisyphodemus will be found throughout the Cockpits in suitable habitat and possibly also in other areas of wet limestone forest (e.g., Mt. Diablo on the border of the Parishes of St. Ann-St. Catherine). Several other species of wet forest frogs ( Hyla marianae, Eleutherodactylus grabhami and possibly E. junori) have disjunct populations on this isolated limestone plateau. Etymology.—The name is Greek, loosely derived from “demos,” district or country, and “Sisyphus,” a king of mythology whose tenure in Hell was spent endlessly rolling a large stone to the top of a hill only to have it roll back down again. As we climbed the fiftieth of what seemed an unending series of steep, crumbling limestone ridges clothed in luxuriant vegetation and inhabited by opaque clouds of mosquitos, we began to gain an appreciation of all things Sisyphean. However, the experience gave us insights into the paradox of why Sisyphus was a happy man. Acknowledgments Jeremy F. Jacobs accompanied me on both forays into the Cockpits and has contributed much to my recent Jamaican field work. Barbara A. Harvey was helpful on the 1975 trip; her monumental patience and good humor often tempered our intolerance and made the trip much more pleasant. George J. and Joan Jacobs also provided indispensable logistical support, enthusiasm, and encouragement. My good friend Menocal Stephenson of Quick Step first introduced me to the Cockpits in 1970. Since then, his assistance and hospitality have been largely responsible for the success of my collecting in the Parishes of Trelawny and St. James. Lester Haggis, also of Quick Step, was a dependable guide and good company on the long walk across the Cockpits. 204 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Albert Schwartz kindly examined the 1974 specimens at my request and confirmed the fact that they represented a new species. George R. Zug, W. Ronald Heyer, and Frances I. McCullough (Smith- sonian Institution) read and commented on the manuscript; Ron Heyer kindly examined the musculature of the new species in the course of his research, and Fran McCullough drew Fig. 1. I am grateful to all for their assistance. My Jamaican field work was partially supported by a grant from the Penrose Fund of the American Philosophical Society. Additional financial support was provided by the Fish and Wildlife Service, through Howard W. Campbell. Literature Cited Asprey, G. F., and R. G. Robbins. 1953. The vegetation of Jamaica. Ecol. Monogr. 23(4) :359-412. Goin, C. J. 1954. Remarks on the evolution of color pattern in the gossei group of the frog genus Eleutherodactylus. Ann. Carnegie Mus. 33(10):185-195. —. 1960. Pattern variation in the frog Eleutherodactylus nubicola Dunn. Bull. Florida State Mus. 5(5) :243-258. Heyer, W. R. 1975. A preliminary analysis of the intergeneric relationships of the frog family Leptodactylidae. Smithson. Contr. Zool. (199 ):1-55. Schwartz, A. 1958. Four new frogs of the genus Eleutherodactylus (Leptodactylidae ) from Cuba. American Mus. Novit. (1873) :1-20. 1969. The Antillean Eleutherodactylus of the auriculatus group. Stud. Fauna Curacao and other Carib. Is. 30(114):99-115. — —. 1971. A new species of bromeliad-inhabiting galliwasp (Sauria: Anguidae ) from Jamaica. Breviora (371 ):1-10. , and D. C. Fowler. 1973. The anura of Jamaica: A status report. Stud. Fauna Curacao and other Carib. Is. 43( 142) :50-142. Starrett, P. H. 1968. The phylogenetic significance of the jaw musculature in anuran amphibians. Unpubl. Ph.D. thesis, Univ. Michigan, viii + 179 pp. Thomas, R. 1975. The argus group of West Indian Sphaerodactylus (Sauria: Gekkonidae). Herpetologica 31(2):177-195. Versey, H. R. 1972. Karst of Jamaica. In Herak, M. and V. T. Springfield (eds.). Karst. Important karst regions of the Northern Hemisphere. Elsevier Pub. Co., Amsterdam, xiv + 551 pp. (pp. 445-466). Division of Reptiles and Amphibians, National Museum of Natural His- tory, Smithsonian Institution, Washington, D.C. 20560. PROC. BIOL. SOC. WASH. 90(2), pp. 205-208 THE SYNONYMY AND DISTRIBUTION OF THE ESTUARINE HYPANIOLA FLORIDA (HARTMAN) FROM THE EAST COAST OF THE UNITED STATES (POLYCHAETA: AMPHARETIDAE ) Marian H. Pettibone In 1953, I described some ampharetids from James Pond, a salt pond on Martha's Vineyard, Massachusetts, under the name of Hypaniola grayji. Some years later, I identified some ampharetids from St. Johns River in Florida as H. grayi. A later review of the literature of the polychaetes from Florida and the Gulf of Mexico has revealed that my species had been briefly described previously by Hartman (1951) under the name of Amphicteis gunneri floridus. It is, thus, belatedly and with regret, that I must now synonymize my species grayi, established in honor of the late Milton (Sam) Gray, my good friend and collector par excellence, with the species H. florida. This synonymy was suggested verbally to Robert Zottoli, then a graduate student at the University of New Hampshire, and his study on the development of this species was published under the name Am- phicteis floridus Hartman (Zottoli, 1974). The purpose of this paper is to record the synonymy and to expand the known distribution of the species, based on previous published records, as well as collections sent to me for identification. The latter material is now deposited in the Smithsonian Institution (USNM). Annotated Synonymy of Hypaniola florida (Hartman), new combination Amphicteis gunneri floridus Hartman, 1951:110 [Englewood, at lower end of Godfrey Creek, emptying into Lemon Bay, western Florida, and Och- lockonee Bay, northwest Florida].—Carpenter, 1956:93, 106 [Alligator Harbor, Franklin County, Bald Point, south side mouth of Ochlockonee Bay, northwest Florida]—Oglesby, 1961:233 [Estuary of Apalachicola River at Apalachicola, Franklin County, northwest Florida; host of Cer- caria hartmanae Oglesby |. Hypaniola grayi Pettibone, 1953:384 [James Pond, Martha’s Vineyard, Massachusetts ].—Stickney, 1959:17, 18 [Sheepscot River Estuary near Wiscasset, Maine].—Jones and Burbanck, 1959:123 [Upper Estuary of Pocasset River, Cape Cod, Massachusetts].—Sanders, Mangelsdort and Hampson, 1965:R222 [Upper Estuary of Pocasset River, Cape Cod, Massachusetts ].—Wall, 1973:682 [West Harwich, Barnstable County, Cape Cod, Massachusetts]|.—Kinner, Maurer and Leathem, 1974:689 206 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON [Delaware Bay].—Williams, Hogan and Zo, 1975:27, 28, 30 [Hudson River, vicinity of Ossining, New York].—Harrel, Ashcraft, Howard and Patterson, 1976:75, 76, 78 [Neches River estuary, Texas]. Amphicteis floridus.—Zottoli, 1974:78 [Upper reaches of Piscataqua River, New Hampshire; observations on early development]. Material examined—NEW HAMPSHIRE: Mouth of Johnson’s Creek, Durham, upper reaches of Piscataqua River, July 1960, May and June 1962, November 1964, M. Pettibone, L. Scott, R. Zottoli and R. Gilmore, collectors —15 specimens (USNM 54037, 54041-2, 54045). MASSACHUSETTS: James Pond, Martha's Vineyard, summer 1944, August 1950, 1951, 1952, M. B. Gray and M. Pettibone, collectors—7 syntypes (USNM 24734-5) and 8 specimens (USNM 54028-9). West Harwich, Barnstable County, Cape Cod, July 1968, W. J. Wall, collector—7 specimens (USNM 54046). Pocas- set River, Cape Cod, December 1957, W. Burbanck, collector—9 specimens (USNM 54032). Pamet River, small creek at extreme end of Cape Cod near Provincetown, October 1957, W. Burbanck, collector—6 specimens (USNM 54031). Stony Brook, Cape Cod, May 1958, W. Burbanck, collector —4 specimens (USNM 54034). CONNECTICUT: Old Mystic on Mystic River, head of estuary, 4°23/15’N, 71°57’43’W, November 1962 and June 1964, J. Rankin and M. Mazurkiewicz, collectors—17+ specimens (USNM 94043-4). DELAWARE: Indian River, Millsboro Dam, east of Millsboro, Sussex County, March 1970, R. D. Jones, collector—2 specimens (USNM 54050). MARYLAND: Salt marsh north of Ocean City, Eastern shore, May 1953, M. Pettibone, collector—28 specimens (USNM 54030). Wiscomico River, Charles County, head of estuary, May 1958, J. P. E. Morrison, col- lector—7 specimens (USNM 54033). Patuxent River at Long Point, 1 mile south of Benedict, August 1968, S. L. H. Fuller, collector—3 specimens (USNM 54047). Salt marsh west of Colton, St. Mary’s County, July 1959, F. W. Grim, collector—50+ specimens (USNM 54035). VIRGINIA: Machodoc Creek, May 1961, M. Wass, collector—25+ specimens (USNM 54039). NORTH CAROLINA: Morton’s Mill Pond, North Harlowe, back- water of Albermarle Sound, summer 1961, J. L. Taylor, collector—12 speci- mens (USNM 54040). Currituck Sound, October 1960, J. A. Kerwin, col- lector—4 specimens (USNM 54038). EASTERN FLORIDA: St. Johns River, Brevard County, February 1955, Wm. McLane et al., collectors — : 1 specimen (USNM 27387). NORTHWESTERN FLORIDA: Estuary of Apalachicola River, July 1960, L. Oglesby, collector—3 specimens (USNM 54036). Keaton’s Beach, Suwannee River, December 1959, L. C. Oglesby, collector—50+ specimens, including many small ones (USNM 39664, | 50474-5). Haven Point, North Bay, Panama City, surface of Diplanthera | flats, October 1957, M. L. Jones, collector—5 specimens (USNM 50473). Woodlawn, St. Andrews Bay, Panama City, October 1957, M. L. Jones, 4 VOLUME 90, NUMBER 2 207 collector—4 specimens (USNM 50472). MISSISSIPPI: Ocean Springs, on Escatawpa River just north of Orange Grove, 30°26’N, 88°27’W, March 1969, W. W. Langley, collector—1l specimen (USNM 54048). LOUISI- ANA: Lake Pontchartrain, July 1975, M. A. Poirrier, collector—3 speci- mens (USNM 54466-7). TEXAS: Upper part of Galveston Bay, about 6 miles above mouth of Cedar Bayou (very polluted tributary of Trinity Bay), December 1969, G. E. Williams, collector—54 specimens (USNM 54049). Fence Lake in Sea Rim State Park, Jefferson County, 13 Septem- ber 1975, B. J. Callahan, collector—4 specimens (USNM 54902). Remarks.—Hypaniola florida is a small ampharetid polychaete (up to 30 mm in length) occupying tubes up to several times its length composed of mucus, sand, mud and debris. It forms a part of the infauna, and is found in salt ponds, marshes, estuaries and river mouths, often in consider- able numbers. At times it may be the dominant animal in the estuary (Sanders, et al., 1965). It is found on bottoms of sand, mud, muddy sand, firm clay, with gravel, plant debris and places rich in bio-detritus. H. florida may be classed as oligo- and mesohaline, being found in waters of low salinity from less than 1% to 30%. It may be found associated with other estuarine polychaetes, such as the nereids Nereis diversicolor O. F. Miller and Laeonereis culveri (Webster ), the glycerid Glycera dibranchiata Ehlers, the orbiniid Scoloplos fragilis (Verrill), the spionids Scolecolepides viridis Verrill, Polydora ligni Webster and Streblospio benedicti Webster, and the capitellid Heteromastus filiformis Claparede. In Johnson's Creek, New Hampshire, the ampharetids were observed being engulfed by the minute mud anemone Nematostella sp. (L. Scott, in litt.). In New Hamp- shire, large yolky eggs were laid in the tubes of the females, from late May to early September, where they were fertilized and developed into non- pelagic larvae. The latter left the adult tubes at the two- to three-setiger stage, when they crawled on the surface of the mud, began feeding and formed mucous tubes (Zottoli, 1974). Distribution—Estuaries along the east coast of the United States from Maine to Florida and Gulf of Mexico (Florida to Texas). Acknowledgment I wish to thank my colleague Dr. Meredith Jones for reviewing the manu- script and for his helpful suggestions. Literature Cited Carpenter, D. G. 1956. Distribution of polychaete annelids in the Alligator Harbor area, Franklin County, Florida. Florida State Univ. Studies, no. 22:S9—110. Harrel, R. C., J. Ashcraft, R. Howard, and L. Patterson. 1976. Stress and com- munity structure of macrobenthos in a Gulf coast riverine estuary. Contr. Mar. Sci. 20:69-81. 208 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Hartman, O. 1951. The littoral marine annelids of the Gulf of Mexico. Publ. Inst. Mar. So. l(t) s7aey Jones, N. S., and W. D. Burbanck. 1959. Almyracuma proximoculi gen. et sp. nov. (Crustacea, Cumacea) from brackish water of Cape Cod, Massachusetts. Biol. Bull. Woods Hole 116(1):115—124. Kinner, P., D. Maurer, and W. Leathem. 1974. Benthic invertebrates in Delaware Bay: Animal-sediment associations of the dominant species. Inter. Rev. Hydrobiol. 59(5) :685-701. Oglesby, L. C. 1961. A new cercaria from an annelid. Joum. Parasitology 47(2): 233-236. Pettibone, M. H. 1953. A new species of polychaete worm of the family Ampharetidae from Massachusetts. Journ. Washington Acad. Sci. 43(11):384-386. Sanders, H. L., P. C. Mangelsdorf, Jr., and G. R. Hampson. 1965. Salinity and faunal distribution in the Pocasset River, Massachusetts. Limnol. Oceanogr. 10 (Suppl.): R216—-R229. Stickney, A. P. 1959. Ecology of the Sheepscot River Estuary. Special Scient. Rep. Fish. No. 309. U.S. Depart. Interior Fish and Wildlife Serv., 21 pp. Wall, W. J., Jr. 1973. The intertidal sand and salt marsh invertebrate fauna as- sociated with the bloodsucking Diptera of Cape Cod, Massachusetts. Environ. Entom. 2(4):681-684. Williams, B. S., T. Hogan, and Z. Zo. 1975. The benthic environment of the Hudson River in the vicinity of Ossining, New York, during 1972 and 1973. New York Fish and Game Journ. 22(1):25-31. Zottoli, R. A. 1974. Reproduction and larval development of the ampharetid poly- chaete Amphicteis floridus. Trans. Amer. Micros. Soc. 93(1):78—89. Department of Invertebrate Zoology, Smithsonian Institution, Washington, D.C. 20560. PROC. BIOL. SOC. WASH. 90(2), pp. 209-213 SCALE ORGANS ON THE HEAD OF LEPTOTYPHLOPS (REPTILIA, SERPENTES): A VARIATIONAL STUDY Braulio Orejas-Miranda, George R. Zug, Daniel Y. E. Garcia, and Federico Achaval Introduction Miniature hairlike filaments, pits, and protuberances have long been recognized as common surface features of squamate scales. Only in snakes have the scale organs been used frequently as taxonomic characters, pri- marily scale pits for the differentiation of species. Concentrations of scale organs have been observed on the heads of scolecophidian snakes (e.g., Haas, 1930; McDowell, 1974; Jackson and Reno, 1975), but have not at- tracted the attention of taxonomists. Using small samples of Leptotyphlops dulcis and L. munoai, we wish to show that the number and distribution of the scale organs on the head scale are of potential value in future taxonomic studies. Materials and Methods A series of nine Leptotyphlops dulcis (Carnegie Museum 52942 A-I; Texas, Bexar Co., San Antonio) and six L. munoai (Museo de Historia Natural de Montevideo 1626; Uruguay, Depto. Rio Negro, Estacion Francia; and Departmento de Zoologia, Facultad de Ciencias, Montevideo R349, R1015-16, R1018, R1023; Uruguay, Depto. Tacuarembo, Pozo Hondo) were examined for the distribution and number of scale organs on their heads. Both left and right sides were counted and recorded. The distribution of scale organs was mapped on outline drawings of the head, similar to Fig. 1. Results and Discussion For most head scales, L. dulcis possesses more scale organs per scale than does L. munoai (see Table 1). This difference is particularly evident for the anterior and anterolateral head scales (rostral, upper nasal, lower nasal, ocular, anterior supralabial, and first infralabial) which are statistically significantly different. Such differences clearly suggest that the number of scale organs on these six scales is of diagnostic value and should be analyzed in future Leptotyphlops taxonomic studies. A high degree of symmetry exists between left and right sides. The homogeneity between the two sides of the head in each snake was tested by chi-square. For Leptotyphlops dulcis, the calculated chi-squares range from 0.304 to 2.574 and in L. munoai from 1.794 to 8.482 (x7,0.05) = 22.4 210 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 1. Diagrammatic illustration of the head scales of Leptotyphlops. A. Lateral view. B. Dorsal view. C. Ventral view. Abbreviations: ASI, Anterior supralabial; F, frontal; II, infralabial; Io, interoccipital; Ip, interparietal; LN, lower nasal; M, mental; O, ocular; Oc, occipital; Pa, parietal; Pf, prefrontal; Po, postoccipital; PS], posterior supralabial; R, rostral; So, supraocular; T, temporal; UN, upper nasal. Terminology follows Klauber (1940:figs. 1-5). for 13 df). These values demonstrate the strong similarity in the number of scale organs on the opposite sides of the head. Even without the histological data (see Landmann, 1976, for literature | review ), the concentration and distribution of the scale organs on the head | indicate a sensory function. The anteriormost scales (rostral, upper and lower nasals) possess the highest number. The scale number shows a gradual decline from anterior to posterior on all surfaces (Fig. 2). Simi- | larly, the absence of these organs from the dorsoanterior quarter of the | ocular scales, the integument precisely external to the eye, indicates a | segregation of sensory structures and functions. The pioneering histological — VOLUME 90, NUMBER 2 211 Table 1. Summary of scale sense organ number variation in Leptotyphlops. The same abbreviations for the scales are used as in Fig. 1. Only the value for the left of paired scales is included. dulcis (n = 9) munoai (n = 6) Scale x Range SD x Range SD R 188.0 NGQ=i, 5a 90.7 44-116 262 Pf Oil 5-7 0.9 By 0-5 1.9 F 2.8 9-5 1.0 0.2 0-1 0.4 Ip KE 2-4 0.7 0 0 0 Io - - - 0 0 0 UN DEO 50-63 4.4 30.7 24-39 6.4 So 2.8 1-4 1.0 CD 4-10 Meso: O 44.1 40-47 2.6 24.8 17-31 4.5 Pa 16.3 13-19 1.6 Tol 4-11 oul Oc 4.8 2-7 Met 1.0 0=2 0.9 LN 38.4 35-43 3.4 alee) 19-26 3D ASI 22.4 20-25 1.9 12.5 11-15 1.4 PSI ISL 11-18 2.8 da 10-14 Wes T 3.0 2-4 0.5 DP 2-3 0.4 mei Dil 13-27 4.5 9.7 5-14 onal 11-2 20.4 17-28 3.3 15.0 13-18 a7 11-3 6.8 6-8 0.8 6.5 3-14 Delt 11-4 2.6 2-6 1.3 2.4 1-5 1 1-5 1.2 1-2 0.4 LS 1-2 0.4 study in scolecophidian snakes (Aota, 1940) and the recent comparative histological survey in reptiles (Landmann, 1976) show the organs clearly to be integumentary sense organs. The function of scale organs was long assumed to be tactile and recently, Hiller (1968) experimentally showed the hairlike scale organs on a gecko’s foot to be mechanoreceptors. It seems likely, therefore, that the majority of integumentary sense organs possess a similar capability. We suspect that for scolecophidian snakes, the organs have a pressure-sensor role, which serves as a guidance mechanism during burrowing. Aside from their function, much remains to be discovered about these sense organs. A thorough survey of their variation in a single species over a wide geographic area is necessary to test their reliability and stability as a taxonomic character. Our samples show relatively low variability at a single locality and suggest, but do not prove, that variation may be low over a large geographic expanse. Jackson (1971) reported intraspecific varia- tion in pattern, numbers, and distribution, but did not contirm this varia- tion in his more detailed report (Jackson and Reno, 1975). If stability or low variation occurs intraspecifically, an interspecific survey should demon- strate whether integumentary sense organ number can be used as a genetic 212 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON DO A SO SIEINSIE ORGANS ant. / eS ee eS. DOS. SCALE F2OSiMNON Fig. 2. The number of scale organs per scale in Leptotyphlops munoai for three anteroposterior transects. A. Sagittal transect: rostral, prefrontal, frontal, interparietal, and interoccipital. B. Parasagittal: upper nasal, ocular, parietal, occipital. C. Lower lip: first infralabial to fifth infralabial. relationship indicator or ecological-behavior equivalence indicator. We | hope these problems will attract other investigators. | Summary The distribution and number of integumentary sense organs per scale on | the head scales are examined in Leptotyphlops dulcis and L. munoai. The § anterior scales bear the greatest number of sense organs, and number de- creases rapidly from the anterior to the posterior head scale. Intraspecific | variation is relatively low, and a high degree of symmetry exists in the | number of organs on paired scales. This relative stability of sense organ | number and the significant difference in scale number on the anterior and | anterolateral scales of the two species recommend these counts as new | taxonomic or diagnostic characters. | VOLUME 90, NUMBER 2 213 Acknowledgments We wish to thank the members of the Division of Reptiles and Am- phibians, Smithsonian Institution, for their constant helpfulness and C. J. McCoy, Carnegie Museum, for use of specimens and friendly advise. Literature Cited Aota, Shigetada. 1940. An histological study on the integument of a blind snake, Typhlops braminus (Daudin), with special reference to the sense organs and nerve ends. Jour. Sci. Hirosima Univ., ser. B. 7( 10) :193-208. Haas, Georg. 1930. Uber das Kopfskelett und die Kaumuskulatur der Typhlopiden und Glauconiiden. Zool. Jahrb., Abt. Anat. Ontog. Tiere 52:1-94. Hiller, U. 1968. Untersuchungen zum Feinbau and zur Funktion der Haftsborsten der Reptilien. Zeits. Morphol. Tiere 62( 4) :307—362. Jackson, Morris K. 1971. Cutaneous sense organs on the heads of some small ground snakes in the genera Leptotyphlops, Tantilla, Sonora, and Virginia (Reptilia: Serpentes). Amer. Zool. 11(4):abst. 732. , and Harley W. Reno. 1975. Comparative skin structure of some fossorial and subfossorial leptotyphlopid and colubrid snakes. Herpetologica 31(3):350—359. Klauber, L. M. 1940. The worm snakes of the genus Leptotyphops in the United States and northern Mexico. Trans. San Diego Soc. Nat. Hist. 9(18):87—162. Landmann, Lukas. 1976. The sense organs in the skin of the head of Squamata. Israel Jour. Zool. (1975), 42:99-135. McDowell, Samuel P. 1974. A catalog of the snakes of New Guinea and the Solomons, with special reference to those in the Bernice P. Bishop Museum, Part 1. Scolecophidia. Jour. Herpetol. 8(1):1—57. (BO-M) Department of Scientific Affairs, Organization of American States, Washington, D.C. 20006; (GRZ) Division of Reptiles and Amphibians, National Museum of Natural History, Washington, D.C. 20560; (DYEG&FA) Museo de Historia Natural de Montevideo, Montevideo, Uruguay. PROC. BIOL. SOC. WASH. 90(2), pp. 214-217 DISTRIBUTIONAL AND NOMENCLATURAL NOTES ON SOME GELASTOCORIDAE IN THE COLLECTION OF THE BERNICE P. BISHOP MUSEUM (HEMIPTERA) ld, Ibs INOIDID A collection of 381 adult gelastocorid bugs, mostly from New Guinea, from the Bernice P. Bishop Museum, Honolulu, Hawaii has been identified as follows. The distributional data, number of specimens of each sex, and the location of the material is presented for the record to facilitate research by future students of the family. These data constitute another contribu- tion to the continuing series of lists and faunal revisions published sub- sequent to the revision of the family; Todd, 1955, Univ. Kansas Sci. Bull., 37, pt. 1(11):277-475. Gelastocoris angulatus (Melin), 1929, Zool. Bidrag fran Uppsala, 12:169. BRAZIL: Para, Belem, 1é, 12. Gelastocoris hungerfordi Melin, 1929, Ibidem, 12:168. GUATEMALA: Escuintla, El Salto, 2é. Gelastocoris vicinus Champion, 1901, Biol. Centri-Amer., Rhynch., Het., 2:347. GUATEMALA: Escuintla, El Salto, 16, 12. Nerthra adspersa (Stal), 1863, Berliner Entomol. Zeitschr., 7:407. AUS- TRALIA: Western Australia, Bluff Knoll, Sterling Ranges, 1¢. Nerthra ampliata (Montandon), 1899, Bull. Soc. Sci. Bucarest, 8(4/5) :404. NEW GUINEA: Papua, Areke Ptn., N. Distr., 2¢; Papua, Carberry Ptn., N Distr., 2¢; Kokoda, 14; Koitaki, 16, 12; SE New Guinea, Popondetta, WD Nerthra cheesmanae Todd, 1959, Nova Guinea, n. ser., 10, pt. 1:822. Nerthra stevensi Todd, 1959, Ibidem, n. ser., 10, pt. 1:87 brachypterous ¢. [New synonymy. ] Size in mm: Males (19), length 9.8 to 11.8, mean 11.0; width of head 4.4 to 5.0, mean 4.8; width of pronotum 7.0 to 8.1, mean 7.7; width of abdomen 6.9 to 7.8, mean 7.8. Females (15), length 10.9 to 12.4, mean 11.7; width of head 4.7 to 5.0, mean 4.85; width of pronotum 7.3 to 8.2, mean 7.9 mm; width of abdomen 7.3 to 8.2, mean 7.7 mm. NEW GUINEA: NE New Guinea, Mt. Kaindi, 16 km SW of Wau, 12¢, 72; NE New Guinea, Mt. Wilhelm, 16, 12; NE New Guinea, 29-32 km S of Wau, Bulldog Rd., 16; NE New Guinea, Morobe Distr., Wau, 4¢, 52; NE New Guinea, 32 km E of Wapenamunda, 1é; Lake Sirunki, 12; IMipnarany 12. ILS. VOLUME 90, NUMBER 2 215 The macropterous males of this species have genitalia that agree exactly with those of the male type of N. stevensi Todd. The latter is now con- sidered to be only an individual brachypterous specimen and a junior synonym of N. cheesmanae Todd. The unique type of cheesmanae was from Papua; therefore the specimens listed and the inclusion of stevensi increase the known range of the species. Nerthra colaticollis Todd, 1959, Ibidem, n. ser., 10, pt. 1:75. NEW GUINEA: Neth. New Guinea, Genjam, 40 km W of Hollandia, 1¢, 22; Neth. New Guinea, Guega, W of Swart Val., 16; Neth. New Guinea, Kutsime, W of Swart Val., 16; W New Guinea, Star Mts., Sibil Val., 4¢, 52; W New Guinea, Hollandia, 12; NW New Guinea, Vogelkop, Sucumi Camp nr. head of Ransiki R., 12; W New Guinea, Vogelkop, Kebar Val., W of Mano- kwari, 32; NW New Guinea, Ifar, Cyclops Mts., 16,32; NW New Guinea, Nabire, S Geelvink Bay, 2¢, 62; NW New Guinea, Moanemani, Kamo V., Wisselmeren, 12; NE New Guinea, 16 km NW Banz, 2°; NE New Guinea, Purosa, E Highlands, 46; NE New Guinea, 13 km. SE Okapa, 36,32; NE New Guinea, Lae, 16, 12; NE New Guinea, Purosa, 20-26 km SE of Okapa, 1¢, 22; NE New Guinea, Wau, Morobe Distr., 26, 9°; NE New Guinea, Dreikikir, Sejik Distr., 12; SE New Guinea, Jumbora, 34, 22; SE New Guinea, Oriomo River, 66, 112; SE New Guinea, above Tigobi nr. Tari, 1¢; Papua, W District, Oriomo Govt. Sta., 14, 1°; Papua, Rigo District, 3¢, 22; Japen I., SSE of Sumberbaka, Dawai R., los LS. Nerthra conabilis Todd, 1959, Ibidem, n. ser., 10, pt. 1:88. NEW GUINEA: SE New Guinea, above Tigobi nr. Tari, 1¢; SE New Guinea, Mt. Gilowe, 6¢, 92; NE New Guinea, Yaibos, W Highlands, 16; NE New Guinea, Simbai, Bismark Range, 246, 22; NE New Guinea, 1 km S of Mt. Hagen (town), 1¢; NE New Guinea, 16 km NW of Banz, 14; NE New Guinea, 6-12 km W of Wabag, 16; NE New Guinea, Laleng, Huon Pen., 12; NE New Guinea, 32 km E of Wapenamunda, 12; NE New Guinea, Tuwep, Salawaket Range, 12; NE New Guinea, Kepilam, 2°; NE New Guinea, Maprik, 12; NE New Guinea, 11 km S of Laiagam, 1°; NE New Guinea, Wau, Morobe Distr., 10¢, 52; NW New Guinea, Enarotadi, Wisselmeren, 9¢, 122; Papua, Anga gorge, E of Mendi, 1°. Nerthra gurneyi Todd, 1955, Univ. Kansas Sci. Bull., 37, pt. 1(11):419. SOLOMON ISLANDS: Fauro I., 12; San Cristoval, Wugiroga, 1¢; San Cristoval, Mensi, 12; San Cristoval, Bweinaniawarikiapu, 12; Santa Ysabel, Sukapisu, 1¢; San Ysabel, Tamatah-Koloau, 1°. Nerthra hirta Todd?, 1959, Nova Guinea, n. ser. 10, pt. 1:83. NEW BRIT- AIN: Gazelle Pen., Upper Warangoi, 12. 216 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON This species was previously known only from New Guinea. The doubtful determination is given because of the sex of the specimen. Nerthra improcera Todd, 1959, Ibidem, n. ser. 10, pt. 1:73. NEW GUINEA: NE New Guinea, Mt. Piora, 1°. Nerthra indica (Atkinson), 1888, Journ. Asiatic Soc. Bengal, 57:345. CHINA: Kwangtung, Chukiang, Lungtaushan, 16, 22. These are the first specimens recorded from China. Nerthra infecta Todd, 1959, Nova Guinea, n. ser., 10, pt. 1:91. NEW GUINEA: NE New Guinea, Bulldog Rd., 14 km S of Edie Ck., 16; NE New Guinea, Korn Farm, W Highlands, 1¢; NE New Guinea, Sinofi, 1¢; NE New Guinea, Tuwep, Salawaket Range, 12; NE New Guinea, Tapo (Tapu), 3 km NW of Kainantu, 12; NW New Guinea, Vogelkop, Surwai, SW of Lake Anggi Giji, 12; Neth. New Guinea, Enarotadi, Wisselmeren, 1¢; Neth. New Guinea, Wamena, 22. Nerthra laticollis (Guérin-Meneville), 1843, Rev. Zool. Travaux Ined., 6: 112. NEW GUINEA: NE New Guinea, Wau, Morobe Distr.,9¢, 32; NW New Guinea, Ifar, Cyclops Mts., 22. NEW BRITAIN: Gaulim, Gazelle Pen., 12; Upper Warangoi, Gazelle Pen., 12; Illugi, Upper Warangoi, Gazelle Pen., 26, 52. Nerthra luteovaria ( Distant), 1904, Ann. Mag. Nat. Hist., (7), 14:63. AUS- TRALIA: Western Australia, Robe River, 1¢; Northern Territory, Kathe- rans, oy ee The example from Robe River is the first recorded from Western Australia. Nerthra macrostyla Todd, 1955, Univ. Kansas Sci. Bull., 37, pt. 1(11):428. SOLOMON ISLANDS: Santa Ysabel, Tamatshi, 1¢; San Ysabel, Horora, 2°; Kolombangara, Iriri, 12; Kolombangara, Pepele, 73, 52; New Georgia Group, Gizo I., 12; Vella Lavella, Ulo Crater, 19. Nerthra macrothorax (Montrouzier ), 1855, Ann. Sci. Phys. Nat. Lyon, 2:110. PHILIPPINE ISLANDS: Leyte, Palo, 1¢. Nerthra mixta (Montandon), 1899, Bull. Soc. Sci. Bucarest, 8(4/5) :404. NEW GUINEA: NE New Guinea, Wau, Morobe Distr., 7¢, 152; Lae, 12; NE New Guinea, Dreikikir, Sepik Distr., 12; Neth. New Guinea, Genjam, 40 km W of Hollandia, 22. NEW BRITAIN: Illugi, Upper Warangoi, Gazelle Pen., 1°. Nerthra mixtella Todd, 1959, Nova Guinea, n. ser., 10, pt. 1:85. NEW GUINEA: NE New Guinea, Purosa, 20-26 km SE Okapa, 26; NE New Guinea, 24-26 km, SE Okapa, 1¢; NE New Guinea, E Highlands, Purosa, 26,12; NE New Guinea, Karimui, 16, 22; NW New Guinea, Enarotadi, Wisselmeren, 16; West New Guinea, Star Mts., Sibil Val, 16, 49; VOLUME 90, NUMBER 2 217 NE New Guinea, 13-20 km SE of Okapa, 12; NE New Guinea, Wau, Morobe Distr., 1°. Nerthra omani Todd, 1955, Univ. Kansas Sci. Bull., 37, pt. 1(11):422. NEW BRITAIN: near Rabaul, 1¢. SOLOMON ISLANDS: San Cristoval, Napagiwae, 18¢, 132; Guadalcanal, Tambalia, 35 km W of Honiara, 14; San Cristoval, Bweinaniawarikiapu to Manpwena, 1?; Nuna Lava, Malaita, 25 km NE of Dala, 12. The male from near Rabaul, New Britain is the first specimen studied from that island. Nerthra recta Todd, 1959, Nova Guinea, n. ser., 10, pt. 1:80. NEW GUINEA: Neth. New Guinea, Genjam, 40 km W of Hollandia, 146, 12; NE New Guinea, Wau, Mt. Missim, Morobe Distr., 1°. The specimen from Wau is doubtfully determined as this species because of the sex. Nerthra robusta Todd, 1955, Univ. Kansas Sci. Bull., 37, pt. 1(11):429. NEW GUINEA: NE New Guinea, Wau, Morobe Distr., 36,52; NE New Guinea, Lae, Singuawa R., 12; Bulolo R., 1°. Nerthra tasmaniensis Todd ?, 1955, Ibidem, 37, pt. 1(11):437. AUS- TRALIA: New South Wales, Mt. Victoria, 1°. Nerthra toxopeusi Todd ?, 1959, Nova Guinea, n. ser., 10, pt. 1:92. PHILIPPINE ISLANDS: Mindanao, Agusan, S Francisco, 1°. Because the specimen is a female, the determination must be considered uncertain, but it appears to be toxopeusi which was previously known from Buru, Ceram, and Halmahera. The only species previously known from the Philippine Islands is the very distinct, widespread species, N. macro- thorax (Montrouzier ). Nerthra unguistyla Todd, 1957, Proc. Ent. Soc. Wash., 59(4):152. INDIA: Karikal, 1é. Falls Church, Virginia, c/o U.S. National Museum, NHB-127, Washington, D.C. 20560. PROC. BIOL. SOC. WASH. 90(2), pp. 218-231 VARGULA TSUJII, A NEW SPECIES OF LUMINESCENT OSTRACODA FROM LOWER AND SOUTHERN CALIFORNIA (MYODOCOPA: CYPRIDININAE) Louis S. Kornicker, and James H. Baker Many species of Ostracoda in the subfamily Cypridininae are luminous, but no species in this subfamily have previously been described from the western coast of North America. The new species described herein, Vargula tsujii, was observed to be luminous by a collector who gave the speci- mens to Professor Frederick I. Tsuji, University of Southern California, a specialist in bioluminescence, who forwarded specimens to the senior author for identification. Additional specimens were supplied by Brad Myers, J. L. Barnard, and the junior author. The new species seems to be fairly abundant in some areas along the coasts of Lower and southern Cali- fornia and therefore, may prove useful as a nearby source of luciferin and luciferase, the glandular secretions known to produce luminescence in Ostracoda. The holotype and some of the paratypes have been deposited at the National Museum of Natural History, Washington, D.C.; paratypes have been deposited at the Allan Hancock Foundation, University of Southern California, Los Angeles. A key to the known species of Vargula is included. Station Data and Numbers of Specimens of Vargula tsujii Santa Catalina Island and Californian shelf; coll. Allan Hancock Founda- tion R/V Velero IV: Sta. 2847; 23 June 1954; 6.3 mi 182° T (True N) from West End Light, Santa Catalina Island; 33°22’30’N, 118°36’38’W; 931 m (1 ad. ¢ ).—Sta. 3208; 7 July 1955; 4.0 mi 262° T from Santa Monica Pier Light, 34°00'03”N, 118°34’40’”W; 45.7 m (1 ovig. 2? ).—Sta. 5201; 15 August 1957; 33°19’15’N, 117°36’30’W; 88.1 m; sediment temperature 10.8°C; orange peel grab (28.3 liters); debris (1).—Sta. 5631; 22 February 1958; 33°22’50’N, 117°34’55’W; 17.1 m; orange peel grab (0.8 liters); coarse gray sand and red algae (1 ad. ¢ ).—Sta. 5745; 16 May 1958; 33°34’47’N, 118°08’17’W;; 83 m; orange peel grab (12.4 liters) (1 ovig. ?).—Sta. 5749; 16 May 1958; 33°36’10”N, 118°15’25’W; 78.6 m; orange peel grab (3.7 liters) (1).— Sta. 6432; 30 September 1959; 2.3 mi 327.5° T from Pinos Point Light; 36°39’57’N, 121°57’30’W; 89.6 m (1).—Sta. 6435; 30 September 1959; 3.65 mi 302° T from Pinos Point Light; 36°39/57’N, 121°59’50’W;; 115.2 m (1). Santa Catalina Island: Big Fishermans Cove, collected about 1975, by Tony Chess; probably epibenthic (8, including 4 ovig. 2 @ and 1 ad. ¢).— Offshore, coll. about 1975, Tony Chess; probably epibenthic (2 ovig. 2 2, VOLUME 90, NUMBER 2 219 3 juv.).—Santa Catalina Marine Biological Station, University of Southern California, plankton net, 1975; received from F. Tsuji (1 ad. 2 ). La Jolla, California, 7 May 1968, water depth 24 m; from body of the pomocentrid fish Chromis atrilobata; coll. E. S. Hobson (1 juv. ¢ ). Bahia Los Angeles, Baja California, Mexico; Sta. SI0-62-216, 21 April 1962, shore sample, reef between Isla Ventana and Isla Cabeza de Caballo, tailings of fish sample; coll. Carl L. Hubbs and party, Scripps Institution of Oceanography; received from J. L. Barnard (1 ovig. 2, 4 ad. 4 4, 9 juv.). Cypridininae Baird, 1850 The senior author takes this opportunity to correct a couplet in a pub- lished key to the genera of Cypridininae (Kornicker, 1975:86). 17. Furcal claws 2 and 4 united with lamella Siphonostra Only furcal claw 2 united with lamella or no furcal claws united with lamella Skogsbergia Vargula Skogsberg, 1920 Type-Species.—Cypridina norvegica Baird, 1860. Key to the Species of Vargula (Expanded from Poulsen (1962:178) and Kornicker (1975:154), but with many changes.) 1. Furca with 12 or more pairs of claws. V. hilgendorfii (Muller ) Furca with 11 or less pairs of claws. 2 2. No furcal claws united with lamella. 3 Only 2nd furcal claw united with lamella. 4 2nd and 4th or 2nd, 3rd and 4th furcal claws united with lamella. 5 3. Tip of 7th limb with dorsal jaw. V?. danae ( Brady ) Tip of 7th limb with dorsal peg. V. sutura Kornicker 4. Tip of 7th limb with dorsal peg. V. tsujii, new species Tip of 7th limb with dorsal jaw. 6 5. Tusks on upper lip with prominent proximal tooth. 7 Tusks on upper lip without proximal tooth. 8 6. Dorsal jaw at tip of 7th limb bare. V. spinosa Poulsen Dorsal jaw at tip of 7th limb with spinelike hairs. V. spinulosa Poulsen 7. Posterior of carapace with tubelike caudal process. V. tubulata Poulsen Posterior of carapace with narrow caudal process. V. dentata Kornicker 8. Seventh limb with hook-shaped process within terminal comb. V. hamata Kornicker 220 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON I ma IPSS Dein. nie TS H F EB = = | E = = = = = = = = Ale | = LJ e S| s' Bas By \3 SI a a is Oo A el A LT . Sr Fig. 1. Vargula tsujii, holotype, ovig. 2, USNM 156767, length 2.12 mm: a, Com- plete specimen showing central adductor muscle attachments, right lateral eye and position of eggs as seen through shell; b, Joints 1-4 and proximal part of 5th joint of right Ist antenna, medial view; c, Joints 6-8 and distal part of 5th joint of right Ist VOLUME 90, NUMBER 2 221 Seventh limb with dorsal jaw opposite terminal comb. V. harveyi Kornicker and King Seventh limb with single peg opposite terminal comb. 9 9. Lateral eyes absent. V. lusca Kornicker Lateral eyes of female with 4 or 5 ommatidia. 10 Lateral eyes of female with 9 or more ommatidia. ml 10. Carapace longer than 2.8 mm; lateral eyes of females about % length of rod-shaped organ. V. antarctica (Miller ) Carapace shorter than 2.25 mm; lateral eyes of females slightly longer than rod-shaped organ. V. stathme Kornicker 11. Height of caudal process about % height of valve. __-V. plicata Poulsen Height of caudal process % to % height of valve. 12 12. Interval between c- and d-bristles of mandibular basale not con- siderably larger than width of larger d-bristle. | V. bullae Poulsen Interval between c- and d-bristles several times larger than width of larger d-bristle. 3 13. Caudal process projecting sufficiently to form well-defined angle where dorsal edge of process joins posterior margin of valve. V. puppis Poulsen Caudal process very narrow, forming little or no angle with posterior margin of valve. 14 14. Lateral eye with 9 or 10 ommatidia. V. norvegica (Baird) Lateral eye with 14-16 ommatidia. V. subantarctica Kornicker Vargula tsujii, new species Figs. 1-6 Etymology.—The species is named for Dr. Frederick I. Tsuji, University of Southern California, who submitted some of the specimens studied. Holotype —USNM 156767, length 2.12 mm, ovig. 2 on slides and in alcohol. Type-locality — Bahia Los Angeles, sta. SIO 62-216. Allotype —USNM 157133, length 2.21 mm, ad. ¢ from same sample as holotype. Paratypes——USNM 157134a, b, 2 ad. 6, same sample as holotype; USNM 157135, 1 ad. ¢ with torn carapace, and 9 juv., same sample as holotype; Baker slide 253, valves of $, USC sta. 2847; Baker slide 254, ad. ¢ ap- < antenna, medial view; d, Endopodite and distal part of protopodite of right 2nd antenna, lateral view; e, Coxale endite and some bristles on ventral margin of basale of right mandible, medial view; f, 3rd endopodial joint and distal part of 2nd endopodial joint of left mandible, medial view. 222 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON pendages, USC sta. 2847; 1 mounted specimen, USC sta. 6435; 1 ovig. 2, USC sta. 3208; 1 specimen, USC sta. 5201; 1 ad. ¢, USC sta. 5631; 1 speci- men, USC sta. 5749; 1 ovig. 2, USC sta. 5745; 1 specimen, USC sta. 6432. Big Fisherman’s Cove, Santa Catalina Island: USNM 1571386A, B, 2 ovig. 2; USNM 157136C, 1 ad. ¢; 2 ovig. 2 plus 3 specimens returned to Brad Myers. Off Santa Catalina Island: 1 ovig. 2 and 1 juv. returned to Brad Myers. USNM 152300, 1 ad. 2 and 1 ad. ¢, off Santa Catalina Island Marine Station; USNM 149195, 1 juv. ¢, off La Jolla, California. Description of adult female (Fig. 1-3a, b).—Carapace (Fig. la) similar in shape to that of adult male. Infold: Bristles not counted, but distribution similar to that of adult ¢; list of caudal process similar to that of adult ¢. Size in mm: USNM 156767, length 2.12, height 1.28; ovig. 2, USC sta. 3208, length 2.31, height 1.39; ovig. 2, USC sta. 5745, length 2.26, height 1.35; USNM 157136A, length 2.10, height 1.17; USNM 157136B, length 2.05, height 1.17; 2 specimens, Big Fisherman’s Cove, Catalina Island, length 2.09, height 1.16, and length 2.03, height 1.16. First antenna (Fig. lb, c): First joint bare; 2nd joint with short spines forming rows; 3rd joint short, with 2 bristles (1 ventral, 1 dorsal) and few short medial spines; 4th joint elongate, with 2 bristles, 1 ventral, 1 dorsal; Sth joint about % length of 4th; sensory bristle of 5th joint with 8 long proximal filaments, 2 shorter and more slender distal filaments, and bifurcate tip; medial bristle of 6th joint short, bare. Seventh joint: a-bristle about same length as bristle of 6th joint, with short marginal spines; b- bristle about % longer than a-bristle, with 5 or 6 short marginal filaments; c-bristle longer than sensory bristle of 5th joint, with 9 marginal filaments (some pectinate) and bifurcate tip. Eighth joint: d- and e-bristles bare, about twice length of b-bristle; f-bristle about % length of c-bristle, with about 5 marginal filaments (some pectinate) and bifurcate tip; g-bristle longer than c-bristle, with 9 marginal filaments (some pectinate) and bifurcate tip. Second antenna: Protopodite with short medial bristle (Fig. 1d). Endop- _ odite 3-jointed (Fig. 1d): First joint with 4 proximal bristles (1 long, 3 _ short) and 1 very long distal bristle; 2nd joint elongate with 1 short distal bristle; 3rd joint short, separated from 2nd joint by weak suture, with long | terminal bristle with blunt tip. (Endopodite illustrated in Fig. 1d bears _ fine hairlike filaments interpreted as a foreign growth; this growth was ob- _ served on many appendages of the specimen.) Exopodite: First joint bare; | bristle of 2nd joint reaching 8th or 9th joint, with 11 stout ventral spines; | natatory bristles without spines; 9th joint with 4 bristles of which short dorsal bristle bears a few short faint spines, remaining long bristles with natatory hairs but no ventral spines; basal spines of joints 3-8 increasing in size on distal joints; basal spine of 8th joint extending past distal end of | nt ev RA — - VOLUME 90, NUMBER 2 223 srt COMUMLHDGE SS WSS Li XS Nie \ LS ANY ANANS ah v7 Ki LSS a Fig. 2. Vargula tsujii, holotype, ovig. 2, USNM 156767: a, Left maxilla, posterior view (exopodite and some bristles of basale not shown); b, Endites and exopodite of right 5th limb, posterior view; c, Right 6th limb, medial view; d, Anterior of body show- ing medial eye, rod-shaped organ, anterior process, and upper lip; e, Right lateral eye, anterior to right. 224 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON G, Mmmmroar 7 eg 4! PHU fy IT 1 Fig. 3. Vargula tsujii, holotype, ovig. 9, USNM 156767: a, 7th limb; b, Tip of 7th limb shown in “a”. Adult ¢, paratype from USC sta. 5631, length 1.73 mm: c, Central adductor muscle attachments of right valve, outside view, anterior to right. Adult ¢, allotype, USNM 157133, length 2.21 mm: d, Complete specimen showing position of VOLUME 90, NUMBER 2 225 9th joint; lateral bristle of 9th joint about same length as basal spine of 8th joint. Mandible (Fig. le, f): Coxale endite well developed, spinous (terminal spine stouter than others); basal bristle well developed. Basale: ventral margin with 2 a-bristles (1 long, 1 very short), 1 very short b-bristle (about same size as short a-bristle), 2 c-bristles (1 short, bare, and 1 long with short marginal spines ), and 2 long d-bristles (proximal of these % to % length of distal bristle and with short marginal spines; remaining bristle with long marginal spines); dorsal margin with 3 bristles (1 subterminal, 2 ter- minal; 1 of terminal bristles about same length as subterminal bristle, other terminal bristle about twice length of other; both terminal bristles with short marginal spines). Exopodite about % the length of dorsal margin of Ist endopodite joint, with 2 subterminal bristles (proximal bristle about twice length of other). Endopodite: First joint with 4 ventral bristles (1 of these minute, 1 short, 2 long); ventral margin of 2nd joint with 2 short, single, ringed bristles, 1 short, ringed, subterminal, medial bristle, and a subterminal, lateral process with rounded tip reaching past distal end of 3rd joint; dorsal margin of 2nd joint with 5 long bristles, 1 medium bristle, and 5 short bristles (1 of these with long stout spines along both margins); medial side of 2nd joint with short faint spines forming short rows near dorsal margin. End joint with total of 7 claws and bristles (a pair of medial bristles near ventral margin, ventral of these minute; a short ventral bristle with bulbous base and ringed, narrow, distal part, possibly with opening at tip; 1 lateral and 1 medial claw near dorsal margin, both slender; 1 middle claw with broad base bearing medial teeth and 1 short, lateral bristle near dorsal margin). Maxilla (Fig. 2a): Coxale with stout plumose bristle and dorsal fringe. Endite I with about 9 spinous bristles; endite II with about 4 spinous ter- minal bristles; endite III with about 6 spinous bristles, 1 proximal, 5 distal. Basale with 2 short bristles along distal margin. Exopodite with 3 bristles of about same length (proximal and posterior, terminal bristle plumose ). Endopodite: First joint with spines along dorsal margin, a terminal tooth on inner margin having 2 teeth, 1 alpha-bristle with few long hairs, and 1 beta-bristle with spines along inner margin; end joint with 3 a-bristles with teeth along anterior margins, 2 b-bristles (anterior of these with spines along posterior margin, other with spines along anterior margin ), 3 c-bristles < right lateral eye; e, Anterior of left valve, inside view; f, Anterior of right valve, outside view; g, Posterior of left valve, inside view; h, Posterior of right valve, outside view; i, Anterior of body showing medial eye, rod-shaped organ, anterior process and upper lip; j, Right lateral eye, anterior to right. Adult ¢@, paratype, Baker slide 254 from USC sta. 2847: k, Left lamella of furca, lateral view; 1, Upper lip viewed from right side. 226 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON (anterior of these minute, remaining 2 with spines along anterior margins), and 3 d-bristles with spines along anterior margins. Fifth limb (Fig. 2b): Epipodial appendage with about 48 bristles. Endite I with 6 spinous bristles; endite II with 5 spinous bristles; endite II with 5 or 6 spinous bristles. Exopodite: main tooth of Ist joint with proximal smooth peg and 6 larger pectinate teeth; a pectinate bristle present near proximal peg of main tooth; 2nd joint with posterior bristle with long proximal hairs, about 13 terminal pectinate bristles, and 1 long spinous anterior bristle; inner lobe of 3rd joint with 1 short bristle with long proximal hairs and 1 long bristle with short, faint, marginal spines; outer lobe of 3rd joint with 2 bristles with short marginal spines; 4th and 5th joints fused, with total of 2 bristles with short marginal spines; joints 3-5 with abundant long hairs. Sixth limb (Fig. 2c): Three short bare bristles in place of epipodial ap- pendage. Endite I small, with 2 short medial bristles and long terminal bristle; endite II with 3 short medial bristles and 2 long terminal bristles; endite III with 1 medial bristle and 3 terminal bristles (middle bristle short); endite IV with 1 medial bristle and 1 terminal bristle. End joint with 6 spinous anterior bristles separated by space from 1 short spinous bristle and 2 longer, plumose, posterior bristles; limb hirsute. Seventh limb (Fig. 3a, b): Comb side with 6 distal and 2 to 4 proximal bristles; peg side with 5 distal and 4 proximal bristles; each bristle with up to 5 bells and without marginal spines; comb with 10-12 short square- tipped teeth (4-6 on each side), and 9 longer alate middle teeth; a single smooth peg present opposite comb. Furca: Same as that of adult male described herein. Medial eye and rod-shaped organ (Fig. 2d): Medial eye pigmented; rod- shaped organ short with more-or-less rounded tip. Lateral eye: Pigmented, about twice diameter of medial eye, with about 19 ommatidia, not all ommatidia shown in illustration (Fig. 2e). Genitalia: Consisting of 2 sclerotized oval rings, each with short lobe extending ventrally; 1 ring on each side of body anterior to furca. Upper lip (Fig. 2d): Similar to that of adult male described herein. Eggs: USNM_ 156767-8; specimen from USC sta. 5745-8; USNM 157136A-8; USNM 157136B-9; specimen from Big Fishermans Bay, Cata- lina Island, returned to Brad Myers—10. Description of adult male (Fig. 3c-l, 4-6 ).—Carapace elongate with con- vex dorsal and ventral margins, prominent caudal process and deep incisur (Fig. 3d-h); anterior margin of rostrum evenly rounded; surface of valves smooth. Infold (Fig. 3e, g): Rostral infold with 10-13 double bristles forming row parallel to anterior margin of rostrum and 1 double bristle proximal to row; 2 bristles present at inner end of incisur; anteroventral and ventral 227 VOLUME 90, NUMBER 2 Joints 1-4 b, Joints 6-8 and distal a Endopodite and distal part of S47: 9 sta. Fig. 4. Vargula tsujii, paratype, Baker slide 254 from USC and proximal part of 5th joint of right Ist antenna, lateral vie part of 5th joint of right Ist antenna, lateral vie protopodite of left 2nd antenna. W, Cy Wy 228 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON infold with 23-26 closely spaced double bristles followed by 6 short widely spaced bristles; infold of caudal process with list having minutely crenulate posterior edge; about 13 minute bristles present anterior to crenulate edge of list; about 10 minute bristles or processes present close to outer edge of caudal process. Central adductor muscle scars: Consisting of 15-19 oval scars (Fig. 3c). Size in mm: USNM 157133, length 2.21, height 1.21; USNM 157134a, length 2.20, height 1.27; USNM 157134b, length 1.66, height 0.90; specimen, USC sta. 5631, length 1.73, height 0.96; USNM 157136C, length 1.73, height 0.97. First antenna (Fig. 4a, b): Joints 14 similar to those of ad. 2; sensory bristle of 5th joint with 9 long proximal filaments, otherwise similar to that of ad. 2; 6th joint similar to that of ad. 2. Seventh joint: a-bristle short, bare; b-bristle with short proximal process with broad rounded base and large distal disc; stem of b-bristle with 2 filaments with minute proximal tooth and 5-7 minute discs; c-bristle with proximal process similar to that on b-bristle but with somewhat larger disc; stem of c-bristle with 2 fila- ments with minute proximal tooth and 5 minute discs, other filaments bare except for terminal spine. Eighth joint: bristles in general similar to those Ohad noe Second antenna (Fig. 4c): Similar to that of ad. &. Mandible (Fig. 5a): Coxale endite similar to that of female, but missing from illustrated limb (Fig. 5a). Basale, exopodite, Ist and 3rd endopodial joints similar to those of adult female; 2nd endopodite joint similar to that of 2 except 2 single bristles of ventral margin closer together on illustrated right limb (Fig. 5a), left limb of that specimen and both limbs of another specimen examined had bristles separated as on joint of °. Maxilla and 5th limbs (Fig. 5b, c): Similar to those of ad. 2. Sixth limb (Fig. 5d): Four short bare bristles in place of epipodial ap- pendage. Endite I with 2 short medial bristles and 2 long terminal bristles; endite II with 3 short medial bristles and 2-4 long terminal bristles; endite III with 1 medium length medial bristle and 3 terminal bristles (middle bristle short); endite IV with 1 medial and 1 terminal bristle. End joint with 5 spinous anterior bristles separated by a space from 1 short bristle with long proximal and short distal spines, and 2 long plumose posterior bristles; limb hirsute. Seventh limb (Fig. 6b): Comb side with 6 or 7 bristles, 4 or 5 distal, 1-3 proximal; peg side with 9 bristles, 4 or 5 distal, 4 or 5 proximal, each bristle with up to 6 bells and without marginal spines; comb and peg similar to thoserot adi 2. Furca (Fig. 3k): Each lamella with 9 claws; claw 2 fused to lamella, remaining claws separated from lamella by suture; claws with 1 or 2 rows VOLUME 90, NUMBER 2 f dS eR Sas as SSN BI y Athy ee La SL \ 4 OY \ 4 L/ a0! G6 A 4 Ss \ \ n0oe Se = // ea KA \ ei) se cS ae \ < fy \| a s) iu U i 0 YS QW RQ =e, SS Swe, SSS asa i) = SS wy, pees! oy ZA ra C1 fh A “fy So / Za soo oot = Z Fig. 5. Vargula tsujii, paratype, Baker slide 254 from USC sta. 2847: a, Right mandible, medial view (not all marginal spines on bristles shown); b, ec, Distal parts of 5th limbs; d, Right 6th limb, medial view. of teeth along posterior margin; claws 14 with faint spines along anterior Margin. Medial eye and rod-shaped organ (Fig. 3i): Medial eye and _ rod- shaped organ similar to that of adult female except rod-shaped organ with triangular tip. 229 230 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON WY a SA Ses Fig. 6. Vargula tsujii, paratype, Baker slide 254 from USC sta. 2847: a, Copulatory appendage; b, 7th limb. Lateral eye: Similar to that of lateral eye of 2, but with about 16 ommatidia, not all ommatidia shown in illustrated eye (Fig. 3j). Upper lip (Fig. 3i, 1): Tip and distal posterior margin of each tusk with a fairly large glandular projection; process at tip longer on some speci- mens than on others; a minute spine may be present at position of distal posterior glandular process. Lip otherwise typical for genus. Copulatory organ (Fig. 6a): Long outer lobe terminating in recurved process with scooplike tip; inner lobe with 10-12 short bristles and ter- minating in knob with minute pustules; a smaller proximal lobe with 2 bristles may be present but is obscure on mounted organ of allotype. Claw- like process on copulatory organ of Macrocypridina castanea (Brady, 1897) (see Kornicker et al., 1976:16) not observed on the present species. Comparisons.—Only 3 previously described species of Vargula have only the 2nd claw united to the lamella without an intervening suture: V. hilgendorfii (Muller, 1890), V. spinosa Poulsen, 1962, and V. spinulosa Poulsen, 1962. V. hilgendorfii has 12 furcal claws compared to only 9 on V. tsujii. The tip of the 7th limb of V. tsujii bears a single peg opposite the comb, whereas, both V. spinosa and V. spinulosa have a stout jaw op- posite the comb. Miller (1890:233) referred specimens collected along the west coast of Central America to a new species, Pyrocypris americana Miller, 1890. In lateral outline the carapace of that species differs from V. tsujii in having the anterior margin of the rostrum forming a definite angle. The anterior margin of the rostrum of V. tsujii is evenly curved. Also, the carapace of VOLUME 90, NUMBER 2 231 P. americana bears 7 hairs along the anteroventral margin that project well past the margin. These hairs are not present on V. tsujii. Miiller referred his species to Pyrocypris Miller, 1890 (=Cypridina Milne-Edwards, 1840), a genus characterized by having an upper lip with 2 single anterior processes and 2 pairs of posterior processes. The upper lip of species of Vargula are without anterior processes and have only 1 pair of posterior processes. We enquired at several European institutions and museums seek- ing Miller's specimens in order to make a direct comparison, but were unsuccessful; the specimens must be presumed lost. Literature Cited Baird, W. 1850. The natural history of the British Entomostraca. London. Ray Society, 364 pages, 36 plates. 1860. Note upon the genus Cypridina Milne-Edwards, with a description of some new species. Proceedings of the Zoological Society of London 28:199-202. Brady, G. S. 1880. Report on the Ostracoda dredged by H.M.S. Challenger during the years 1873-1876. Report on the Scientific Results of the Voyage of H.M.S. Challenger, Zoology, 1(3):1-184, pls. 1-44. —. 1897. A supplementary report on the crustaceans of the group Myodocopa obtained during the Challenger Expeditions with notes on other new or imperfectly known species. Transactions of the Zoological Society of London 14(3) (7):85-100. Kornicker, L. S. 1975. Antarctic Ostracoda (Myodocopina). Smithsonian Contribu- tions to Zoology 163:1—720, 432 figures, 9 plates. , S. Wirsing, and M. McManus. 1976. Biological studies of the Bermuda Ocean Acre: Planktonic Ostracoda. Smithsonian Contributions to Zoology 223: 1-34, 20 figures. Milne-Edwards, H. 1840. Ordre des Cyproides ou des Ostracodes. Histoire Naturelle des Crustacés 3:393-411. Paris. Miller, G. W. 1890. Neue Cypridiniden. Zoologische Jahrbiicher 5:211-252. 1908. Die Ostracoden. Deutsche Siidpolar-Expedition 1901-1903, 10(2): 52-178. Poulsen, E. M. 1962. Ostracoda-Myodocopa, 1: Cypridiniformes-Cypridinidae. Dana- Report 57:1—414, 181 figures. Copenhagen (Carlsberg Foundation). Skogsberg, T. 1920. Studies on Marine Ostracods, 1: Cypridinids, Halocyprids and Polycopids. Zoologiska Bidrag fran Uppsala, supplement 1:1—784, 153 figures. Department of Invertebrate Zoology, Smithsonian Institution, Washing- ton, D.C. 20560, and Southwest Research Institute, Houston, Texas 77006. PROC. BIOL. SOC. WASH. 90(2), pp. 232-236 A NEW SPECIES OF APHODIUS (COLEOPTERA: SCARABAEIDAE) FROM SAND DUNES IN CHIHUAHUA, MEXICO Robert D. Gordon In recent years much has been done on the ecology of sand dune areas in western North America. The insect fauna of some of these dunes has proven to be unexpectedly rich with a high percentage of endemics, many of which have not been described. The dunes from which the species of Aphodius described here was taken have not been extensively collected; they may prove to possess an extensive beetle fauna. This large, distinctive Aphodius species has not been recorded from any other locality which may indicate that it is endemic to these particular dunes, and, if so, it is probable that other endemic species exist there. The purpose of this paper is to provide a name that can be used in further sand dune studies. Aphodius giulianii, new species Figs. 1-4 Holotype.—Male, length 5.82 mm, greatest width 2.78 mm. Form elongate, elytra parallel sided, body somewhat dorsoventrally flattened (Fig. 1). Pale brownish yellow; head, pronotum, sutural interval of elytron, tibiae and tarsi dark yellowish red. Head smooth, shiny, lacking any trace of tubercles, fimbriate with long hairs from clypeal angle to eye, surface faintly aluta- ceous, finely punctured, punctures separated by 1 to 3 times a diameter; anterior margin of clypeus feebly emarginate medially, slightly reflexed, anterolateral angle broadly rounded. Pronotum broad, robust, surface smooth, shiny, faintly alutaceous, punctures on disc fine, separated by less than to 3 times a diameter, punctures becoming coarser and denser toward lateral margin; anterolateral angle rounded, lateral border feebly arcuate, distinctly margined, fringed with hair longer than scutellum, posterolateral angle broadly rounded, posterior border margined, fringed with hair shorter than scutellum. Elytron dull, strongly alutaceous, pubescent, pubescence very short on disc, becoming longer and denser toward lateral and posterior margins; lateral border fimbriate, hairs longer than scutellum; intervals flat, feebly, indistinctly punctured; striae weakly impressed, flat bottomed. Ventral surface smooth, shiny, metasternum punctate medially with oblique row of setae bearing punctures on each side of middle. Anterior tibia with apical spur laterally flattened, abruptly narrowed to apex, apex bent inward (Fig. 2); middle tibia with inner spur nearly straight, apex truncate with sharp tooth at inner angle (Fig. 3); apex of hind tibia fringed with long, unequal spines, apical spurs long, straight, slender, inner spur longer than VOLUME 90, NUMBER 2 233 Fig. 1. Habitus of Aphodius giulianii. first tarsal segment, outer spur shorter than first segment. Anterior tarsus with first segment very short, 4 subsequent segments progressively longer, last segment as long as segments 3 and 4 combined; middle and hind tarsi with basal segment as long as segments 2 and 3 combined, segments 2 and 3 subequal, segment 4 short, segment 5 as long as 3 and 4 combined. Genitalia as in Fig. 4. Allotype—Female, length 5.41 mm, greatest width 2.49 mm. Similar to male except pronotum reduced in width, surface dull, strongly alutaceous;: spurs on front and middle tibiae slender, unmoditied. Type-material—Holotype, Mexico, Chihuahua, sand dunes 48 km §S Juarez, 28-IV-1974, Derham Giuliani (USNM 73883). Allotype, same data as holotype (USNM). Paratypes, 8, same data as holotype, in collections of USNM and Alan Hardy, California Department of Agriculture, Sacramento. Variation —Length ranges from 5.0 to 6.0 mm, width from 2.35 to 2.81 mm. The surface of the pronotum varies from shiny to dull depending on the amount of alutaceous sculpture present. Remarks: The Mexican species of Aphodius have not been taxonomically treated since they were included in the Biologia Centrali-Americana by 234 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Figs. 2-4. Aphodius giulianii. 2, Front tibial spur; 3, Middle tibial spurs; 4, Male genitalia. Bates (1887). I have examined representatives of all the described species of Mexican Aphodius and can state that the affinities of giulianii are not with those species but with a group known from the United States and Canada. This is Horn’s (1887) Group L which is mostly restricted to North America from the Great Plains to the Atlantic Coast. A. giulianii most nearly resembles walshii Horn and femoralis Say, but both of the latter 2 species have black, or nearly black, heads and a dark VOLUME 90, NUMBER 2 235 median area on the pronotum. In addition, the elytral striae are deeply impressed on both species and the male tibial spurs are not at all like those of giulianii. I have modified Horn’s key to the species of Group L to include giulianii. Aphodius tenuistriatus Horn will probably be removed from the group when the entire genus is studied. The species is named for Derham Giuliani in recognition of the prodigious amount of beetle collecting he has done in sand dune areas. Key to species of Horn’s Group L 1. Dorsum entirely rusty red; male pronotum not enlarged. tenuistriatus Horn Dorsum not entirely rusty red; male pronotum usually enlarged. 2 2. Head and pronotum entirely black, elytra red. rubripennis Horn Color pattern not as above. 3 3. Lateral margin of pronotum fimbriate with long hairs. 4 Lateral margin of pronotum lacking hairs, or if hairs present, short and indistinct. 3 4. Head and pronotum dark brown to black, at least medially. walshii Horn Head and pronotum pale brownish yellow throughout. — giulianii, n. sp. d. Elytron entirely pale except sutural stria dark, lacking pubescence. 6 Elytron pale with darker, nebulous areas present, distinctly pubescent. 7 6. Punctures on pronotal disc sparse, separated by 2 to 3 times a diameter. subtruncatus LeConte Punctures on pronotal disc dense, separated by less than a diameter. rehni Robinson 7. Disc and anterior % of pronotum densely punctured; elytron with narrow basal and sutural margins pale yellow. femoralis Say Disc and anterior % of pronotum impunctate or with only a few scattered punctures; elytron yellow except large, irregular darkened area on disc, often extending to lateral border. —prodromus ( Brahm ) Knowledge of the habits and habitats of members of this group is incom- plete. A. prodromus is the only imported European species in the group and is a general surface dung feeder most common in bovine dung. A. tenuistriatus is not infrequently collected in bovine dung in the south- western United States, but that may not be its preferred habitat. I have collected large numbers of A. walshii in sheep dung in sand dune areas of North Dakota. A. femoralis is apparently a general surface dung feeder most often collected in the dung of horses and cattle. A. rubripennis is usually restricted to deer dung but will feed on sheep droppings or even horse manure on occasion. A. subtruncatus and rehni are rarely collected, 236 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON and their habits are not known. I have taken one specimen of sub- truncatus in cow dung and another in a prairie dog burrow, but I con- sider these collections accidental and the preferred habitat unknown. Both subtruncatus and rehni are probably detritus feeders in the upper soil layer. The habits of giulianii are also unknown, and I suspect that it is also a detritus feeder restricted to sandy areas. Literature Cited Bates, H. W. 1886-1890. Biologia Centrali-Americana, Insecta, Coleoptera. Pectini- cornia and Lamellicornia. Vol. II, Pt. 2, pp. 1-432. Hom, G. H. 1887. A monograph of the Aphodiini inhabiting the United States. Trans. Amer. Entomol. Soc. 14:1—110. Systematic Entomology Laboratory, IIBIII, Agr. Res. Serv., USDA, U.S. National Museum, Washington, D.C. 20560. PROG) BIOL?) SOC. WASEL 90(2), pp. 237-242 STUDIES OF TROPICAL AMERICAN LEGUMINOSAE-VIII Richard S. Cowan In studies of some recent Brazilian collections, material of several new taxa came to light. In addition, several collections were received which extend our knowledge of the morphology and distribution of a genus I described several years ago. Macrolobium prancei Cowan, sp. nov. Description.—Arbor 3-6 m alta, ramulis glabris, nigris; stipulae per- sistentes, 10 mm longae, 0.4 mm latae, subulatae, ciliolatae; folia glabra, paripinnata, petioli 4-5 mm longi, canaliculati vel breviter alati supra, rhachibus 28-40 mm longis, alatis, aliis ca. 0.6 mm latis; foliola bijugata, sessilia, chartacea, jugo inferiore minore, 6.0-9.5 cm longo, 2-3 cm lato, jugo superiore 13-16 cm longo, 3.5-5.5 cm lato, anguste elliptica ad elliptica, base inequilaterali, angustato, plus minusve acuto, ad apicem acuminata, costa salienti, venulis obscuris; inflorescentiae 4-10 cm longae, ramuligerae, pedunculo 0-5 mm longo, minute puberulo, axe minute puberulo, bracteis deciduis, lanceolatis, 2 mm longis et 0.8-1 mm latis, pedicellis 4-6 mm longis, minute puberulis; alabastra truncato-cordulata ad basem; bracteolae ubique puberulae vel solum ad basem extus, oblongae, acuminatae, S—10 mm longae, 3.5-5 mm latae; hypanthium glabrum, inaequilateraliter cupu- latum, sepala reflexa, glabra, 2 adaxilibus minoribus, plus minusve liberis, 1.5-2 mm longis, 0.7-1 mm latis, triangularibus vel triangulari-lanceolatis, acutis, sepalis caeteris 3-5.5 mm longis, 1.5-2 mm latis, oblongo-lanceolatis, acuminatis; petalum album vel rubrum, oblanceolatum, 9-10 mm longum et 4-5 mm latum, glabrum, lamina rotundata, unguiculo 4-5 mm longo; filamenta basaliter pilosula, 22-25 mm longa; gynoecium rubrum, stigmate capitulato, stylo 22-25 mm longo, glabro, ovario elliptico, ca. 2 mm longo et 1 mm lato, marginaliter leviter puberulo, gynophoro glabro versus apicem, apicem ipsem sparse puberulo excepto, 2 mm longo; furctus non visus. Type-Collection—D. G. Campbell, J. C. Ongley, and J. F. Ramos P-21961 (holotype US, sheet no. 2801087, isotypes A, CUIABA, F, INPA, Kk, MG, MO, NY, P, R, S, U), “Rio Cuieiras, 50 km upstream, near farm of Sr. Nemerio,” Amazonas, Brazil, 8 April 1974. Additional Specimen Examined.—Rio Cuieiras, 2 km downstream trom farm of Sr. Caldez, Amazonas, Brazil, 5 April 1974, D. G. Campbell, J. C. Ongley, and J. F. Ramos 21862 (INPA, Kk, NY, US). Duplicates of these collections were distributed to the herbaria indicated betore I had seen 238 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON them. Consequently, data from them are not included in the preceding description. Distribution—Known only from two collections made in primary forest on terra firme along the same Amazonian river. Discussion.—The closest relationship of M. prancei apparently is with M. guianense (Aubl.) Pulle, but it is not a very close one. The new species differs from it and all others by the shape of the regularly bijugate leaflets and the ramuligerous inflorescences. It is named for G. T. Prance, Director of the scientific program at the New York Botanical Garden and enthu- siastic authority on the flora of Amazonia. Macrolobium limbatum (Poepp. & Endl.) R. S. Cowan variety propinquum R. S. Cowan, var. nov. Description—Arbuscula 4 m alta, ramulis glabris, stipulis non visis; petiolus 8-9 mm longus, micro-puberulus, laminis coriaceis, 32-33.5 cm longis, 12-13 cm latis, equilateraliter attenuato-acutis basaliter, ad apicem longo-acuminatis, ad basem costae micro-puberulis infra, allioqui glabris; inflorescentiae axillares ad apicem ramulorum, axe 4.5-5 cm longo, micro- puberulo, bracteis deciduiis, ca. 1 mm longis et latis, triangularibus, cilio- latis, micro-puberulis extus, intus glabris, pedicellis 4-5 mm longis, minute puberulis, bracteolis 8 mm longis et 3.5 mm latis, obovato-ellipticis, abrupte acutis, minute puberulis extus, intus glabris; hypanthium infundibuliforme, glabrum, 2 mm longum, stipite 1 mm longo, glabro; sepala 4, oblonga, acuta, 6-6.5 mm longa, 2.5-3 mm lata, glabra; petali lamina destructa non visa, unguiculo 5 mm longo, sparse minute puberulo extus, intus villoso; filamenta 15-16 mm longa, sparse pilosa ad basem, antheris 3.6 mm longis et 1.8 mm latis, ovalibus, thecis minute aculeatis; gynoecium fere glabrum, stigmate capitellato, stylo 11-12.5 mm longo, glabro, ovario 3 mm longo et 1 mm lato, sparse minute puberulo adaxialiter ad adaxilem marginem, gyno- phoro glabro, 1.5 mm longo. Type-Collection.—E. Lleras, W. C. Steward, et al. P-17322 (holotype US, sheet no. 2801085, isotypes INPA, NY, K, U), “Behind Sao Paulo de Olivenga,” Rio Solim6es, Amazonas, Brazil, 16 August 1973. Isotypes were not seen because they were distributed before my work was completed. Discussion—This new variety is quite near to Macrolobium limbatum (Poepp. & Endl.) R. S. Cowan var. mucronatum R. S. Cowan, both phyto- geographically and morphologically. The differences separating them are several, but the most obvious is the nearly glabrous gynoecia of var. propinquum (in allusion to the close relationships of the two varieties). In addition, the petioles are shorter in the new variety; its leaflets are larger and equilateral; the pedicels are longer; and the filaments are shorter and almost glabrous but the anthers are larger. The type localities VOLUME 90, NUMBER 2 239 of the two varieties are separated by several hundred miles, but both are in northwestern Amazonia. Subsequent collections in the region may erase the distinctions, for both are known only from their type collections. Dicymbe puncticulosa Cowan, sp. nov. Description.—Arbor 4 m alta, ramuli juvenales leviter pilosulosi, mox glabrescentes, lenticellis salientibus et nigris; stipulae caducae, non visae; petioli 13-16 mm longi, teretes, sparse pilosulosi, rhachibus 5-6.5 cm longis, sparse pilosulosis et nigro-lenticellatis infra, lenticellis salientibus; foliola 4-jugata, petiolulis 3.5-4 mm longis, pilosulosis, teretibus, laminis tenuiter coriaceis, 44-52 mm longis, 28-36 mm latis, ovalibus vel ellipticis, rotundatis vel leviter cordatis, inaequilateralibus ad basem, abrupte acutis ad apicem, glabris supra, infra puncticulatis, costa leviter pilosulosa et salienti; in- florescentiae axillares et terminales, 6-10 cm longae, composito-racemosae, axe pilosuloso, bracteis caducis, non visis, pedicellis 5-7 mm longis, pilo- sulosis, alabastris oblanceolatis, sericeis et cum pilis longioribus erectis; bracteolae oblongae, leviter tomentulosae intus, 11-12 mm longae (glande ad apicem excepto), 3-3.5 mm latae, glande apicali stipitato, stipite 0.5 mm longo; hypanthium ca. 6 mm longum, leviter arcuatum, glabrum, parte apicali discreta; sepala dilute rosea, dissimilia, uno adaxilare maiore, oblongo, 7.5-9 mm longo, 3 mm lato, sparse pilosuloso extus, caeteris sepalis minoribus, plus minusve fusiformibus, 7-9 mm longis, 1.5-2.5 mm latis, glabris; petala 5, sed solum una cum lamina completa, petalo adaxiali cum stipite 5-6.5 mm longo, pilosuloso intus ad basem et ad costam extus, lamina rotundata, 10.5-11.5 mm longa, 7 mm lata, lutea, caeteris petalis glabris, dilute roseis, fusiformibus, acutis, saepe stipitatis, stipite 2-3.5 mm longo, lamina 5-9.5 mm longa, 1-3 mm lata; filamenta 15 mm longa, pilosa versus basem, ad basem breve connata, antheris 1.5 mm longis et 0.8 mm latis; stigma peltato-capitata, stylo glabro, tenui, 17 mm longo, ovario dense pilosuloso, anguste oblongo, 4.5-5 mm longo, 1.5 mm lato, parte libero gynophori ca. 1 mm longo, dense pilosuloso; fructus immaturus 2-3- seminatus, anguste falcato-oblongus, exalatus sed leviter marginatus adaxia- liter, 10.5-11 cm longus, 2.5 cm latus, micro-puberulus marginaliter; semina immatura. Type-Collection.—C. C. Berg, F. A. Bisby, et al P-18158 (holotype US, sheet no. 2801086, isotypes A, CUIABA, F, INPA, K, M, MG, MO, NY, P, R, S, U), “Manaus-Caracarai Road, km 140. Campina on white sandstone,” Amazonas, Brazil, 27 Sept 1973. Additional Collection—Km 130 Manaus-Caracarai Highway, Amazonas, Brazil, 17 Feb 1974, W. C. Steward, J. F. Ramos, and O. P. Monteiro P-20319 (NY, US). Both collections are from the same area, both growing on white sands from the weathering of sandstones upstream. Although the location 240 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON of duplicates of these collections is cited, they were distributed before I had the opportunity to see them. Consequently data from them were not included in the description. Discussion.—The nearest related species, based on morphological criteria, is D. neblinensis R. S. Cowan, but (1) the leaflets of D. puncticulosa are four-jugate instead of in two or three pairs and they are much closer to- gether on the rachis; (2) the new species has regularly spaced minute punctae on the undersides of the leaflets (hence the specific epithet); (3) the pubescence of many parts in D. puncticulosa consists of short hair- lets appressed to the structure-surface, plus much longer erect hairs inter- mixed; (4) the four sepals of the new taxon are larger than the five described for D. neblinensis; and (5) the dimensions of petals and of stamens are significantly different. Although the two species seem well- separated altitudinally (the new one is a lowland “campina” species and the related one is from the slope forest of Cerro de la Neblina at 650- 700 m), they both occur on soils derived from sandstones. This is the fifth species of Section Eremopetala but, unfortunately, the first with flowers having only four sepals. In all other respects, it is easily assignable to this section. Candolleodendron brachystachyum (DC.) R. S. Cowan Discussion.—When I described this genus (Rhodora 68:429-432. 1968), there were very few collections known to me and, unfortunately, I did not request a loan from the Herbier du Centre ORSTOM de Cayenne, French Guiana. Recently I received a collection from there for identification, and in sending back the determination of the specimen as this species I re- marked that from French Guiana I had only seen the De Candolle type at Geneve. This resulted in a shipment of a substantial number of speci- mens by J. J. de Granville, Director of the Department of Botany of ORSTOM, all of which were correctly identified as this species and most of which had been collected since 1968. This note is to present a more complete description, based on a much larger sample of the population in French Guiana. Only organs whose description was inadequate or er- roneous are mentioned. Infrequently, reproductively mature individuals are shrubby and only 0.5-2 m tall, but generally the habit is a small tree about 3 m tall, [(1-2) 3-5(7) m] and with a trunk 5-17 cm in diameter. The bark peels off the branchlets in thin, papery strips. The imparipinnate leaves have two or sometimes four pairs of opposite to subopposite leaflets, borne on petioles up to 10.5 cm long, a rachis as much as 22 cm long, and petiolules (3-) 5-6(-8) mm long. The form of the leaflet blades is regularly elliptic, but the base ranges from rounded-obtuse to acute and the apex from acute to long-acuminate. VOLUME 90, NUMBER 2 241 Fig. 1. Fruit with seeds of Candolleodendron brachystachyum (Oldeman T.858). The inflorescences are borne only on the trunk and larger branches, the axes ranging from 6-11.5 cm long. The yellow, glabrous petal consists of a claw 1.5-2 mm long and a strongly concave blade 15-19 mm long and 10-13 mm wide with the margin irregularly undulate. The numerous, uniform stamens have filaments up to 4 mm long, borne on a fleshy cushion-like torus, from the center of which arises the gyncecium, the ovary containing a large number of ovules in two YOWS. One completely mature fruit has been seen and it was larger than the submature ones described earlier—6.5 cm long and 2.5 ecm wide. The valves are relatively thin, not more than a millimeter thick at any point. They enclose in a closely packed random manner (Fig. 1) a large number of 242 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON seeds, each on an elongate, slender funiculus up to 27 mm long. The funiculus expands abruptly at the seed in a fleshy aril, perhaps one-tenth the mass of the seed. The seed coat (dried) is brown with many vein-like lines extending in a more or less parallel fashion from the hilar region. Called by the indigenous people “boco,” “bocco blanc,” “bocco rouge,” “coucimi,’ and “nemba,” this small tree is found commonly in French Guiana in the eastern half of the country from Cayenne south and west to the Brazilian borders. Whether this reflects the true phytogeography of the plant or the lack of collecting in the western parts is not known to me. It has most often been recorded on hills at 40 m to 155 m elevation in rain forest. It reaches the height of flowering in December; fruiting occurs in March, although full maturity is not achieved until June. The following collections are known, in addition to those cited in my original description of the genus, all from French Guiana: Mont Mahury, Ile de Cayenne, de Granville B.C. 53; Sinnamary R., west of Saut Aristide, de Granville 192; Yaraupi R., foot of Saut Tainoua, de Granville 359, 374; summit of Mont Gauthiot, Yaraupi R. above Saut Polissoirs, de Granville 416; mouth of Grand Tamouri, Saut Bambaye, de Granville 2116; Sail, de Granville B.2485; Arataie R., Oldeman B-549; Yaraupi R., Saut des Polissoirs, Oldeman T-562; Oyapock R., Maripa Trail, Oldeman T-858; Montagne Tigre, Ile de Cayenne, Oldeman 1060; “Layou Mariaflor,” Oyapock, Olde- man 1861; Comté R., Crique Bagot, Oldeman B2023; Approuague, Crique Tortue, Oldeman 2297; Aratage, “téte du saut Parare,” Oldeman 2925; Saut des Polissoirs, Yaraupi R., Oldeman B-3058; Tracé Crique Limonade, Olde- man 3185; Sail, Tracé Belvédere, Oldeman B.4204; Inini R., Station de l'Institut Pasteur, Sastre 1361. All these collections are represented at ORSTOM, French Guiana; presumably they are also at Paris, but I have not seen other duplicates. Smithsonian Institution, Washington, D.C. 20560. PROC. BIOL. SOC. WASH. 90(2), pp. 243-254 THE GECKOES (SAURIA, GEKKONIDAE) OF THE GENUS SPHAERODACTYLUS OF THE DOMINICAN PENINSULA DE BARAHONA, HISPANIOLA Albert Schwartz The Peninsula de Barahona in the Reptblica Dominicana on the Antillean island of Hispaniola lies to the south of the Sierra de Baoruco (and the extreme eastern portion of the mainly Haitian Massif de la Selle) and extends maximally some 50 km to the south of these ranges. Since the peninsula lies in the rain-shadow of the mountains to the north, it is xeric. However, it is not a uniform desert, as a limestone ridge, reaching an elevation of 322 m, more or less bisects it from north to south and forms a barrier or a specialized habitat for several species of amphibians and reptiles. The Sierra de Baoruco, the easternmost of the three massifs which form the spine of the Hispaniolan south island (sensu Williams, 1961), is generally a mesic range. Since this massif reaches the coast of the Peninsula de Barahona in the east and carries mesic conditions to sea level in many places between Barahona and Enriquillo, the apparently montane (but more appropriately mesic) fauna descends to low levels in the extreme east. To the immediate north of the Sierra de Baoruco lies the very xeric (in places below sea level) Valle de Neiba, and to the west the very high and mesic Massif de la Selle in Haiti. Xerophilic species on the peninsula are, therefore, effectively cut off from their relatives to the north by both the intervening high mountains and their concomitant mesic conditions. This Barahona faunule has come to be regarded as a result of the so-called Barahona Entrapment, in that the fauna of the Peninsula de Barahona is distinctive when compared with that of more northern xeric regions on Hispaniola. A major taxonomic problem in this area concerns lizards of the genus Sphaerodactylus. Shreve (1968:5) described a new subspecies of S. notatus Baird from the town of Oviedo south of the mountains, but Schwartz and Thomas (1975:149) considered S. n. randi Shreve to be a subspecies of S. difficilis Barbour. The latter is the most widespread species of Sphaero- dactylus on the Hispaniolan north island, at least in the Republica Domin- icana; its distribution in Haiti is limited to the northwestern littoral (in- cluding Ile de la Tortue) and the Plateau Central at Hinche. The latter authors’ interpretation of the affinities of S. n. randi is due to the fact that, on the south island, a population of S. difficilis occurring from the vicinity of the city of Barahona along the eastern coast of the peninsula (Fig. 1) seemed to form a continuum geographically with the known distribution of S. n. randi. Although local S. difficilis differ somewhat from long series 244 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 1. The Peninsula de Barahona, Reptblica Dominicana, Hispaniola, showing the known distributions of geckoes of the genus Sphaerodactylus. The dot-and-dash line on the left is the Dominico-Haitian border; the dashed lines delimit the provinces of Pedernales (which includes most of the peninsula), Barahona (to the upper right), and Independencia (upper left). The Massif de la Selle and the Sierra de Baoruco extend diagonally from upper left to about 18° latitude in the east. Symbols are: triangles, S. randi; circles, S. difficilis; solid squares, S. armstrongi; hexagons, S. streptophorus; semi-solid squares, S. thompsoni. Symbols for S. difficilis and S. arm- strongi in the upper right lie north of the Sierra de Baoruco. Other species of Sphaerodactylus which occur on the northern slopes of the Sierra de Baoruco and in the Valle de Neiba (S. rhabdotus, S. altavelensis, S. cryphius, S. leucaster) are not shown on the map although they occur within the limits here displayed. of topotypical randi in scutellation, these two populations differ strikingly in pattern details. In addition to S. difficilis (including randi), three other species of Sphaerodactylus are present on the Peninsula de Barahona. One of these (S. thompsoni Schwartz and Franz) is known from only two specimens taken west of the north-south limestone ridge; the affinities of this species are with the otherwise exclusively Haitian S. copei Steindachner. S. arm- strongi Noble and Hassler is a mesophile occurring in the Massif de la Selle and the Sierra de Baoruco. There it is the only species at moderate to high VOLUME 90, NUMBER 2 245 elevations (to 1,775 m) but in the eastern coastal region of the peninsula, S. armstrongi occurs at sea level in mesic situations (river valleys and ra- vines). The species also occurs on the northern slopes of the Sierra de Baoruco south of Cabral. S. streptophorus Thomas and Schwartz is a small species which occurs in the lowland xeric portions of the peninsula but also ascends both the Sierra de Baoruco and the Massif de la Selle to moderate and high elevations (400 m to 1,600 m); this species also occurs in the Vallée de Trouin and the area near Jacmel in Haiti, some 90 km to the west. On the eastern coast of the peninsula, S. difficilis occurs at and near sea level but maybe encountered as high as 370 m (6 km N Enriquillo ). The ecological relationships of these four species are complex. In the simplest terms, S. armstrongi is in the mesic uplands and S. difficilis occurs in the lowlands; but under special circumstances (penetration into the lowlands of mesic upland conditions and flora) the two species may be found sympatrically. Although S. streptophorus also seems to be primarily a xerophile, it ascends the mountains to 1,775 m but has not been taken sympatrically with S. armstrongi. Its lowland range overlaps broadly with that of S. difficilis, and the two species occur syntopically in the region south of Enriquillo. S. thompsoni is not known to be sympatric with any other Sphaerodactylus. Considering its known altitudinal distribution, it might be expected to be sympatric with S. difficilis and S. streptophorus, but not with S. armstrongi. However, the latter possibility cannot be dismissed as the proper approximation of precise ecological situations might well allow this xerophilic species to occur with the mesophilic S. armstrongi. Finally, examination of Fig. 1 indicates that no species of Sphaerodactylus has as yet been taken across the central portion of the peninsula, a distance of some 35 km. It is in this precise region that the relatively high limestone ridge occurs. As rocks and other cover are abundant, it is amazing that no geckoes have been taken from this seemingly suitable region. However, collecting is difficult there, since the flora (Acacia, cacti) is hostile, and eroded and loose limestone offers havens to small lizards which are virtually inaccessible to the collector. Absence of human habitation and its as- sociated debris, as well as scarcity of fallen and decaying arborescent vegetation minimize the chance of finding these diminutive lizards in this area. In August 1975, Michael H. Strahm and I collected on Cayo Pisaje, an islet lying off the eastern shore of the Peninsula de Barahona south- east of the village of Juancho. The islet is small (ca. 1 X 1.5 km), covered with a dense stand of a tall grass, and with scattered coconut palms and a fringe of mangroves and sea grape (Coccoloba). We quickly encountered geckoes which were moderately abundant under the dead Cocos fronds. These nondescript geckoes are one of the most drab members of the genus 246 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON oe mie? for ae uancho (@@ d) YN Fig. 2. Eastern Peninsula de Barahona, showing details of the distribution of S. difficilis (circles) and S. randi (triangles). In the Enriquillo-Juancho-Oviedo (nuevo) region of contact between the two species. The three lower left triangles are the known distribution of S. r. randi; the two center-right triangles are the dis- tribution of S. r. strahmi. in the Antilles. In the short series secured, some individuals showed enough remnant markings of randi to indicate that they were a derivative of that taxon. Since the nearest known randi locality (the type-locality ) lies some 12 km to the southwest, and since we had already taken S. difficilis near Enriquillo (10 km to the northeast), and since it seemed unlikely that a randi-derivative would occur only on Cayo Pisaje, we searched for these lizards on the mainland. A long series of similarly pat- terned and colored lizards was taken in an extensive Cocos grove op- posite Cayo Pisaje. In this grove the lizards were exceptionally abundant in piles of Cocos fronds and old decaying husks. No other lizards were found there. Since S. difficilis was known from localities only 10 km to the north, it seemed possible that intergrades between the two taxa might be found in the intervening area. But collections from the area between Juancho and the coast opposite Cayo Pisaje, as well as between Enriquillo and the Pisaje region were clearly identical with S. difficilis which occurs along the eastern margin of the Peninsula de Barahona. Even within 1 km of the Cayo Pisaje area we secured S. difficilis in Cocos trash, in both coastal and slightly inland situations (Fig. 2); the companion of S. dif- ficilis in such situations was S. streptophorus. Thus, in Cocos trash piles VOLUME 90, NUMBER 2 247 between Enriquillo and Juancho one finds either S. difficilis or lizards re- lated to randi from further south. The distances between these two popula- tions has been narrowed to less than 1 km; it seems unlikely that randi is to be considered as a subspecies of S. difficilis. I therefore consider S. randi to be a species distinct from S. difficilis, a species limited to the Barahona Entrapment. S. randi is known to occur in three separate regions: the type-locality and vicinity; the town of Pedernales on the Dominico-Haitian border; and on Cayo Pisaje and the adjacent mainland southeast of Juancho, all xeric and at sea level. The species has not been found on even the lower montane slopes. The three samples of S. randi differ from each other in coloration and pattern. No specimens have been collected from areas between these localities. Between Juancho and Oviedo, for instance, the road passes away from the coast, and easily-collected sites (such as Cocos groves) do not occur. Between Oviedo and Pedernales there appears to be ample suitable habitat for a xerophile, yet we have never secured S. randi (nor for that matter any Sphaerodactylus with the exception of S. thompsoni) between a locality 3 km NW Oviedo and Pedernales, a distance of 45 km. In interpreting differences between the three populations of S. randi, I propose the descriptions of two new subspecies. The original description (Shreve, 1968:5) was based upon eight specimens. I have examined the holotype, and there are now 180 additional specimens from the vicinity of the type-locality, on 103 of which I have made scale counts and mea- surements. Sphaerodactylus randi Shreve, new combination Definition—A species of Sphaerodactylus with large, acute, strongly keeled, flattened, imbricate dorsal scales, axilla to groin 27 to 39; no area of middorsal granules or granular scales; dorsal body scales with 1 to 5 hair-bearing organs, each with 1 (very rarely 2) hair, around apex. Dorsal scales of tail keeled, acute, imbricate, and flat-lying; ventral scales of tail smooth, rounded, not enlarged midventrally; gular scales usually smooth but occasionally keeled or partially keeled; chest scales smooth; ventral scales rounded, imbricate, axilla to groin 27-37, smooth; scales around midbody 45-57; internasals 0-3 (mode 1); upper labials to mid- eye 3 (rarely 4); escutcheon with a broad and compact central area and extensions to about midpoint of thigh or a little beyond (4-7 < 12-26). Dor- sal coloration and pattern variable, not strongly sexually dichromatic; dorsal ground color from pale tan to dark brown, either uniform or flecked with darker to give a “salt and pepper’ effect; females of one population at times with orange body ocelli; males with heads heavily dotted to vermicu- late dark brown or black on a yellowish or tan ground, heads in females 248 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON either dotted with dark brown or with a trilineate pattern; throat either concolor with venter (flesh, whitish, dark brown) or with upper side of head (yellow); scapular patch (in juveniles) more or less square and black, with a widely opened pale chevron along its anterior face and with two ocelli, either included in or lying along the posterior edge of the scapular patch, these pale ocelli whitish to orange, and at times fused to each other to give a black patch with both anterior and posterior margins pale or orange; shoulder pattern much reduced in males (usu- ally absent but ocelli and bar at times indicated) and less reduced in adult females, which have the patch, bar, and ocelli present but obscure or even lacking. Sphaerodactylus randi methorius, new subspecies Holotype—CM 60501, an adult male, from Pedernales, Pedernales Province, Republica Dominicana, one of two taken 28 June 1964 by D. C. Leber. Original number ASFS V2662. Paratypes—ASFS V2663, same data as holotype; ASFS V2592-93, same locality as holotype, 26 June 1964, R. Thomas; ASFS V2921, same lo- cality as holotype, hatched 9 July 1954 from egg taken 2 July 1964, R. Thomas; ASFS V29855-57, Pedernales, south of center of town, Pedernales Province, Republica Dominicana, 11 August 1971, A. Schwartz. Definition—A subspecies of S. randi characterized by the combination of pale tan dorsal ground color, dorsum flecked or not with slightly darker brown, scapular patch faint to absent in males, indicated to present and an elongate bar in females, anterior pale chevron and posterior pale ocelli present or indicated in both sexes; head trilineate in females, and vaguely trilineate, the pattern overlain by dark spotting, in males; throat finely dotted or stippled dark brown in both sexes; juveniles with prom- inent scapular patch and anterior bar and paired posterior ocelli included within the patch rather than lying along its posterior margin; venter whitish to flesh; throat scales smooth. Description of holotype—An adult male with a snout—vent length of 27 mm and tail 26 mm; dorsal scales between axilla and groin 31, ventrals between axilla and groin 32; midbody scales 51; 3/3 supralabials to eye- center; 1 internasal; 10 fourth toe subdigital lamellae; throat, chest and ventral scales smooth; escutcheon 5 X 14. Dorsum tan with faint darker brown “salt and pepper” speckling; head vaguely spotted darker brown over a faint indication of female trilineate pattern; dark scapular patch very tiny but anterior bar and posterior ocelli present, very faint and pale; throat yellowish with some scattered brown dotting; venter light grayish; underside of tail yellowish. Variation.—The series of S. r. methorius consists of 3 males, 2 females, VOLUME 90, NUMBER 2 249 and 3 juveniles. The largest male (ASFS V29855) has a snout—vent length of 29 mm, the largest female (ASFS V29857) 31 mm. The juveniles have snout—vent lengths of 15 mm and 16 mm; the hatchling has a snout—vent length of 15 mm. Scale counts on the short series are: dorsals 27-31 (x = 29.6), ventrals 30-34 (31.8), midbody scales 50-57 (52.0); all have 3/3 supralabials and 1 internasal; fourth toe lamellae vary between 10 and 13 (M, = 11), and throat, chin, and ventral scales are smooth in all. The escutcheon has 5-6 X 14-23 scales (x = 5.3 x 19.7). Males are tan dorsally, flecked or not with brown; the scapular patch is faint to absent and the anterior bar and posterior ocelli are only in- dicated; the head is dull yellow, dotted with dark brown, but in the 3 males there is still clearly an indication of the trilineate female pattern; the throat is yellowish, flecked with dark brown; the venter is whitish to flesh. Females are colored dorsally like the males but both are “salt and pepper ; the patch is present but elongate and bar-like in one female, and almost absent in the other; bar and ocelli are clearly indicated, the ocelli peripheral and posterior to the patch; the head pattern is trilineate, with a median dark line from the snout to the transverse prepatch bar and two broad lateral lines that extend to above the forelimb insertion and thus are lateral to the patch but not confluent with it. The throat is con- color flesh with the venter and is finely flecked with dark brown. The three juveniles show the basic S. randi pattern most vividly. The head is boldly trilineate, the patch is large, dark, bordered anteriorly by a pale bar and with the two pale ocelli enclosed within the posterior margin of the patch. The name methorius is from the Greek meaning “bordering on,” in reference to the geographic position of Pedernales. Comparisons.—There is some difficulty in comparing the short series of S. r. methorius with the very long series of S. r. randi. The latter ap- pears to reach a slightly larger size (males to 31 mm, females to 32 mm) than S. r. methorius. Scale counts (dorsals, ventrals) average more (32.6, 32.2) in S. r. randi than they do in S. r. methorius (29.6, 31.8), but in both cases, the extremes in S. r. randi encompass those of S. r. methorius. Mid- body scales on the other hand average less (50.6) in S. r. randi than in S. r. methorius (52.0); extremes in the former embrace the range in the latter. All S. r. methorius have 3/3 supralabials to eye-center; this is also the mode in S. r. randi, but counts of 3/4 (11 specimens) and 4/4 (1) also occur. Internasals in S. r. randi are distributed modally as 1 in 9S individuals, but this scale is absent in 1, 2 in 1], and 3 in 1 lizard. Fourth toe lamellae vary between 8 and 14, but the mode (11) is the same in S. r. randi as in S. r. methorius. The escutcheon is slightly longer and broader in S. r. randi with counts of 4-7 X 12-26 (* = 5.8 X 20.3); once again extremes in S. r. randi encompass those in S. r. methorius. In sum- 250 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON mary, scutellar and size differences between the two subspecies seem slight. In color and pattern, however, the two subspecies differ. The dorsal ground color in S. r. randi is gray to dark brown, with a strong tendency toward the darker coloration. The dorsum in both sexes may or may not be flecked with darker scales to give a “salt and pepper” effect. Juveniles have both patch and anterior bar and ocelli present and well developed, but the ocelli lie on the margin of the patch in S. r. randi, not included within it as they are in S. r. methorius. In almost all (57) male S. r. randi, the head shows no indication of the trilineate female head pattern; only one male (ASFS V23205—snout—vent length 28 mm) shows a trilineate pattern. In males the head has a faintly yellow wash and is finely to coarsely covered with dark brown to black dots or blotches. In females, the head is trilineate or covered with dark spotting as in the males. The throat in males is yellowish with dark dotting of the same intensity as that on the top of the head. The throat is concolor with the venter and either unmarked or finely stippled with dark brown in the females. The venter is yellow-gray and the iris is yellow. The scapular patch is absent in males, but vaguely indicated in three of 57 specimens; the patch-associated bar and ocelli (which may be fused to form a posterior bar) are usually also absent, but vaguely indicated in five males. In females, the scapular patch is absent or reduced to form a dark anterior-to-posterior bar; the pre-patch bar and posterior ocelli are always present and are whitish to buffy. In summary, S. r. methorius differs from S. r. randi as follows: pale dorsal color (tan versus gray to dark brown); juveniles with ocelli included within the dark scapular patch rather than lying along its posterior border; and pre-patch bar and ocelli present in males. Remarks.—S. r. methorius has been collected within the town of Peder- nales, about 1 km S of the center of the settlement. The collecting site is an isolated and fairly large shaded Acacia grove that includes scattered Cocos with fallen fronds and also much debris from human habitation. In this grove, S. r. methorius occurs syntopically with S. streptophorus, which is much the commoner sphaerodactyl there. No S. randi were secured when this grove was visited in 1975, but S. streptophorus was more abundant than on previous visits. No Sphaerodactylus have been found elsewhere in the immediate environs of Pedernales and S. randi was taken there only in 1964 and 1971, despite other visits to the Acacia grove. The egg from which the hatchling emerged was taken from a communal nest about 1.4 m above the ground underneath the bark of a standing tree. Since Pedernales lies on the Dominico-Haitian border, separated at this point by the small and intermittent Rio Pedernales, it is probable that S. r. methorius occurs in extreme southeastern Haiti. At the type-locality of S. r. randi, much the same situation prevails except VOLUME 90, NUMBER 2 251 that the more eastern subspecies is locally very abundant. The old town of Oviedo was virtually demolished by Hurricane Inez in 1966. An abun- dance of fallen thatch roofs, boards, and other hurricane wreckage still present in 1969 made collecting S. r. randi a simple task. Visits to the Oviedo area since 1963 have yielded no specimens in undisturbed forest and scrub, nor since 1969 from natural situations despite the abundance of rocks and other debris in the now hurricane damaged forest. The long series from 3 km NW Oviedo (nuevo) was collected by an isolated group of families who took these lizards from debris around human habitations. Undoubtedly S. r. randi is scattered throughout this general region but the population must be minimal in undisturbed situations and thrives only in the immediate vicinity of settlements. Sphaerodactylus randi strahmi, new subspecies Holotype —USNM 197316, an adult male, from 3 km SE Juancho, oppo- site Cayo Pisaje, Pedernales Province, Reptblica Dominicana, one of a series taken 17 August 1975 by A. Schwartz and M. H. Strahm. Original num- ber ASFS V42429. Paratypes.—ASFS V42428, ASFS V42430-33, ASFS V42441-45, ASFS V42460-66, CM 60502-09, MCZ 132363-69, USNM 197317-22, same data as holotype; ASFS V42418-27, Cayo Pisaje, Pedernales Province, Re- publica Dominicana, 15 August 1975, A. Schwartz, M. H. Strahm. Definition—A subspecies of S. randi characterized by the following: dark brown dorsal ground color without darker scales (not “salt and pep- per’); females often with scattered orange ocelli; scapular patch, anterior bar and ocelli much reduced to absent in both sexes; heads of males with tan ground color densely covered with black dots or spots, finely stippled with dark brown in females, never an indication of the trilineate head pat- tern in either sex or in juveniles; venter dull flesh to brown; juveniles with a small dusky gray scapular patch with concomitant bar and ocelli in- dicated; throat scales usually keeled or at least with some keeling, but occasionally smooth. Description of holotype —An adult male with snout—vent length 28 mm and tail length 24 mm; dorsal scales between axilla and groin 39; ventral scales between axilla and groin 30; midbody scales 50; 3/3 supralabials to eye-center; 1 internasal; 11 fourth toe subdigital lamellae; throat scales keeled, chest and ventral scales smooth; escutcheon 6 X 22. Dorsum dark brown and uniform (without darker scales to give a “salt and pepper” effect), head tan dotted or almost vermiculate with black, this pattern ex- tending onto the throat which has a grayish ground; scapular patch and associated bar and ocelli absent; venter dark brown. Variation.—The series of S. r. strahmi consists of 17 males, 22 females, 252 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON and 10 juveniles with snout—vent lengths 13-23 mm. Snout—vent lengths 29 mm in largest males (ASFS V42432, ASFS V42418) and 31 mm in largest female (ASFS V42441). Scale counts for the series are: dorsals 29-39 (x = 32.9), ventrals 28-36 (31.7), midbody scales 47-55 (49.6); supralabials modally 3/3 (44 individuals) with occasional counts of 3/4 (4) and 4/4 (1); internasals 1 (48) or 2 (1); fourth toe lamellae vary between 9 and 12 (M, = 11); throat scales keeled in 19 adults, partially keeled in 9, completely smooth in 6 adults (all juveniles have smooth throat scales). Escutcheon scales 4-7 xX 12-23 (* = 5.8 x 18.6). Males uniform dark brown dorsally, females vary between light and dark brown; neither sex with any darker scales or “salt and pepper’ dor- sum. Some females with scattered and well defined orange ocelli over the back. Dark scapular patch absent in both sexes. Orange anterior bar and paired scapular ocelli present in only 2 males and 8 females. Head tan above, in males, heavily dotted or vermiculate with black; head in females concolor with dorsum (brown) and finely stippled with brown; this same fine stippling continues on throat. Venter varies from dull grayish flesh to dark brown, iris golden. Specimens from the mainland and from Cayo Pisaje were similar in scutellar details, coloration, and pattern. Comparisons.—Size differences between S. r. strahmi and the two other subspecies are minimal; S. r. strahmi is perhaps the smallest. The mean of dorsal scales (32.9) is greater than those of the two other sub- species (29.6, 32.6), but the mean of ventral scales (31.7) lies below those of S. r. methorius (31.8) and S. r. randi (32.2). Midbody scales on S. r. strahmi average less (49.6) than those in S. r. methorius (52.0) and S. r randi (50.6). In all these scale counts, differences are small. S. r. strahmi is distinctive in the large number of individuals (25) with at least some keeling on the gular scales; no S. r. methorius has the gular scales keeled, and only 6 of 112 S. r. randi have the gular scales keeled. S. r. strahmi is a nondescript sphaerodactyl. Were it not for the dusky scapular patch areas in juveniles and the presence of the pre-patch bar and ocelli in a few adults, its relationships would be uncertain. The drab aspect of S. r. strahmi contrasts strongly with the more boldly marked S. r. randi. The subspecies is named in honor of Michael H. Strahm who helped collect many of the type-series. Remarks.—Specimens of S. r. strahmi from Cayo Pisaje were secured on sand under Cocos fronds and under a log. Those from the type- locality were taken in piles of Cocos fronds and decayed husks on sand, invariably from those nearest to the ground. None were found in upper parts of the piles. Although depth and circumference of frond piles affects the occurrence of many Antillean geckoes (i.e., small or shallow piles yield few to no geckoes, whereas deep and large piles which are cooler and VOLUME 80, NUMBER 2 253 more moist yield more specimens), S. r. strahmi was encountered with equal frequency in small, shallow or large, deep piles. The 39 lizards were taken from 5 piles of varying sizes, and it appeared that additional piles would have provided a much larger number. Discussion Evidence for elevating S. randi to a species distinct from S. difficilis depends primarily on distributional patterns of the two taxa in the re- gion between Enriquillo and Juancho. As the drabbest subspecies of S. randi, S. r. strahmi is a virtually patternless dark lizard. The eastern Peninsula de Barahona population of S. difficilis is, on the other hand, pale (tan) with a prominent and unreduced black scapular patch and two white ocelli in females, and a pair of pale ocelli and dark scapular patch occasionally indicated in males. The two species are quite distinctive as far as color and dorsal pattern are concerned. As far as scutellation is concerned, dorsals in the local S. difficilis vary between 24 and 33, whereas those of S. r. strahmi vary between 29 and 39; some overlap is obvious, but the scales in S. difficilis are larger. Rarely (8 of 80 individuals) are the gular scales of S. difficilis keeled, whereas this is the mode in S. r. strahmi. Finally, median subcaudal scales are broader than long in S. difficilis and are not enlarged in S. randi. Geographically, the two species occur very close to each other (Fig. 2). The southernmost record for S. difficilis is 0.6 km E Juancho, whereas the northernmost record for S. randi is the mainland opposite Cayo Pisaje. The distance between these two points is only 0.8 km airline. S. difficilis also occurs at a locality 7 km SW and 1 km E Enriquillo; this locality is only 0.7 km northeast of Cayo Pisaje and slightly further from the only mainland station of S. r. strahmi. In this region, S. difficilis and S. randi seem to be precise ecological equivalents. At the two southernmost stations for S. difficilis, the lizards were taken in Cocos groves, twice on an earthern substrate, and once on a sandy substrate near the ocean. On both Cayo Pisaje and the mainland, the S. r. strahmi were collected on sandy substrate. The area between the closest approximation of the two species is primarily mangroove swamp along the coast itself and salt marsh for some distance inland, situations altogether inhospitable for geckoes. It is possible that some- where in this immediate region S. randi and S. difficilis occur syntopically, but this has not been verified. One problem is that Cocos groves, the sim- plest place for searching for Sphaerodactylus, are few and scattered in this region. Since the road north of Enriquillo extends to moderate eleva- tions on the southern slopes of the Sierra de Baoruco, one has an excellent panoramic view of the entire region between Enriquillo and Oviedo, and 254 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Cocos groves stand out boldly in contrast to other less arborescent vege- tation. It is likely that we have collected in all coconut groves in this area. Acknowledgments Some specimens used in this study were collected under the auspices of National Science Foundation grants G-7977 and B-023603. I have had the capable assistance in the field of Danny C. Fowler, David C. Leber, James A. Rodgers, Jr., Bruce R. Sheplan, James B. Strong, Michael H. Strahm, and Richard Thomas. Specimens have been deposited in the Albert Schwartz Field Series (ASFS), the Carnegie Museum (CM), the Museum of Comparative Zoology (MCZ), and the National Museum of Natural History (USNM). Through the courtesy of Ernest E. Williams, I have examined the holotype of S. n. randi. Literature Cited Schwartz, A., and R. Thomas. 1975. A check-list of West Indian amphibians and reptiles. Carnegie Mus. Nat. Hist. Spec. Publ. 1:216 pp. Shreve, B. 1968. The notatus group of Sphaerodactylus (Sauria, Gekkonidae) in Hispaniola. Breviora, Mus. Comp. Zool., 280:1-28. Williams, E. E. 1961. Notes on Hispaniolan herpetology. 3. The evolution and re- lationships of the Anolis semilineatus group. Breviora, Mus. Comp. Zool., 136: 1-8. Miami-Dade Community College North, Miami, Florida 33167. PROC. BIOL. SOC. WASH. 90(2), pp. 255-262 EUDRILIN EARTHWORMS, INCLUDING A NEW GENUS, FROM EASTERN NIGERIA A. ©; Segun As only two species of eudrilid (subfamily Eudrilinae) earthworms have previously been described and two others recorded from the eastern region of Nigeria, collections were made from Imo, Anambra and Cross River States (three of the four states constituting the old eastern region of Nigeria). Michaelsen (1913) recorded Eudrilus eugeniae from Old Cala- bar and Segun (1976) described a new genus and species, Agrotoreutus nyongii from Nsukka. Segun (In press) also described a new species, Irido- drilus tonyii from Oron and Ekwagbe near Nsukka, and recorded I. roseus from Oron. Earthworms were collected from gardens and cultivated land by dig- ging and hand sorting during the rains of July, August and September, 1976. In the material collected, nine species of eudrilin earthworms in- cluding a species new to science were identified. The new species exhibits a combination of characters which necessitates the erection of a new genus. This paper contains a description of the new genus and species, and records of eight other species belonging to six genera, Agrotoreutus, Ephyriodrilus, Eudrilus, Hyperiodrilus, Iridodrilus and Keffia. 1. Agrotoreutus nyongii Segun, 1976 Agrotoreutus nyongii Segun, 1976:389-393. Type-locality, Nsukka, Anam- bra State. Material—Botanica! and Zoological Gardens, University of Nigeria, Nsukka, June and July 1976; 30 clitellate, 25 aclitellate specimens. 2. Ephyriodrilus afroccidentalis Sims, 1971 Ephyriodrilus afroccidentalis Sims, 1971:533-536. Type-locality, Ibadan, Nigeria. Material—Botanical Gardens, University of Nigeria, Nsukka, June 1976; 3 clitellate specimens. Farmland near market place, Umuagwo, Imo State, August 1976; 40 clitellate and 3 aclitellate specimens. Edge of cassava plantation, Abachebe, Egbema, Imo State, August, 1976; 1 clitellate speci- men. Farmland, Calabar, Cross River State, September 1976; 1 clitellate specimen. Remarks.—The present series matches the description of this species, but one of the specimens collected from Umuagwo has its porophore on 256 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Mouth. ere a: A Soy <4 Sper mathecal Si Ss pore on XIII Ovo - spermathecal pore on XIV XVIII Male pore with penial seta on XVII Fig. 1. A, Hyperiodrilus euryaulos, external features, ventral view; B, Nsukkadrilus mbae, external features, ventral view. segment XVII. This results in the extreme shortness of the seminal groove linking it with the male pore. 3. Eudrilus eugeniae (Kinberg, 1866) Lumbricus eugeniae Kinberg, 1866:98. Type-locality, St. Helena. Material—Botanical Gardens and Green House, University of Nigeria, Nsukka, June and July 1976; 10 clitellate and 6 aclitellate specimens. Farm- land, Calabar, Cross River State, September 1976; 9 clitellate and 5 aclitel- late specimens. 4. Hyperiodrilus africanus Beddard, 1891 Hyperiodrilus africanus Beddard, 1891:236. Type-locality, Lagos, Nigeria. | VOLUME 90, NUMBER 2 257 Material—Botanical and Zoological Gardens, University of Nigeria, Nsukka, June and July 1976; 12 clitellate specimens. Edge of cassava plantation, Abachebe, Egbema, Imo State, August 1976; 2 clitellate speci- mens. Farmland, Ula-Ehudia on Ahoada-Omoku road, Imo State, August 1976; 2 clitellate specimens. 5. Hyperiodrilus euryaulos Clausen, 1967 Figure 1A Hyperiodrilus euryaulos Clausen, 1967:186-189. Type-locality, Guinea savanna, Togo. Material—Edge of cassava plantation, Abachebe, Egbema, Imo State, August 1976; 3 clitellate specimens. Remarks.—Members of the series from Egbema match the description of this species which had previously been reported from Warri, Nigeria. The external diagnostic features are shown in Fig. 1A. 6. Iridodrilus tonyii Segun (In press) Iridodrilus tonyii Segun, In press. Type-locality, Oron and Ekwagbe, Ni- geria. | Material—Farmland near market place, Umuagwo, near Elele, Imo State, August 1976; 1 clitellate specimen. Farmland, Calabar, Cross River State, September 1976; 19 clitellate and 1 aclitellate specimens. 7. Iridodrilus vomiensis Segun (In press ) Iridodrilus vomiensis Segun, In press. Type-locality, Vom and Jos, Benue Plateau State, Nigeria. Material—Farmland, Ula-Ehudia on Ahoada-Omoku road, Imo State, August 1976; 31 clitellate and 3 aclitellate specimens. Edge of cassava plantation, Abachebe, Egbema, Imo State, August 1976; 9 clitellate and 1 aclitellate specimens. 8. Keffia nigeriensis Clausen, 1963 Keffia nigeriensis Clausen, 1963:311-315. Type-locality, Vom, Benue Pla- teau State, Nigeria. Material.——Botanical and Zoological Gardens, University of Nigeria, Nsukka, Anambra State, June and July 1976; 28 clitellate and 10 aclitellate specimens. 258 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 9. Nsukkadrilus, new genus Definition —Eudrilinae with ventral setae fairly widely, and lateral setae more closely paired. Penial setae present. Dorsal pores absent. Male pores paired on raised papillae on segment XVII; ovo-spermathecal pores paired on XIV. Six intestinal gizzards present in segments XX-XXV, esophageal gizzards and supraintestinal glands absent. Unpaired, ventral esophageal sacs present in segments X and XI, paired dorsolateral esopha- geal glands in segment XII. Holandric, testes enclosed in paired testis sacs. Ovaries paired and enclosed in ovarian sacs in segment XIII, each ovary leading by a narrow ovarian duct into lateral wall of receptaculum seminis. Spermathecal atria paired, left and right parts completely separate. Oviduct leads into spermatheca with no opening of its own. Distribution —Nsukka, Anambra State, Eastern Nigeria. Type-species.—Nsukkadrilus mbae, new species. Remarks.—The genus has affinities with Malodrilus—Eudrilus—Hippopera group. It resembles these genera in the fusion of the female and spermathe- cal pores. The fused ovo-spermathecal openings are paired and they are found anterior to the male pores in all these genera. The morphology of the spermathecal system and its relationship with the female system of this new genus is similar to that of Eudrilus. The morphology of the gut differs in both genera and penial setae are only present in Nsukkadrilus. Although in both Eudrilus and Nsukkadrilus paired dorsolateral esophageal glands are present in segment XII and unpaired ventral esophageal sacs occur in X and XI, the former genus has an esophageal gizzard while in the latter only intestinal gizzards are present. Both Malodrilus and Hippopera with esophageal gizzards also lack intestinal gizzards. Nsukkadrilus resembles Hyperiodrilus, Iridodrilus and Heliodrilus in possessing intestinal gizzards, but both female and spermathecal openings are separate in these genera. Of these genera, only Hippopera possesses paired prostatic pores. Etymology.—Nsukkadrilus is named after the University town of Nsukka in Anambra State, Nigeria from where the specimens were collected. Nsukkadrilus mbae, new species Fig, 1B, 2 Material—One of the four clitellate specimens collected from Botanical and Zoological Gardens, University of Nigeria, Nsukka is now kept in the British Museum (Natural History), Reg. No. 1976. 24.1 as the holo- type. One of the three paratypes is also kept in the British Museum (Natural History), Reg. No. 1976.24.2 and the other two in the Natural History Museum, University of Ife, Ile-Ife, Nigeria, Reg. No. 1976.1.1-2. | VOLUME 90, NUMBER 2 259 ee Be ee ite er emai d | {i Ea Ovarian Se af aN vesicle / Press BE f a) OVISAC Samay ay tf 4 Ovo oS Be B y oo Spermathecal- a duct. Vasa- —— deferentia : Euprostate- ( ais he ot gland- wy 4 ne Muscul@je ——4 j sheath of - . HE penial seta. i % : EB £ Ventral- — : — cd; the setal formula (AA:AB:BC:CD) at segment XX = 8:6:8:3 and at segment XXX = 13:5: 4:1. dd = % circumference. The male pores are paired on the apices of 2 protuberances between 2 a setae on segment XVII, both protuberances being joined by a slender 260 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON bridge (Fig. 2). Penial setae are sometimes seen protruding from the male pores. The ovo-spermathecal pores with tumid lips are slightly more closely paired than the male pores on segment XIV (Fig. 2). Genital papillae and markings are lacking. The nephridiopores are paired in the posterior wall of each furrow within setal lines cd, equidistant from c and d. Internal characters.—First septum 4/5 is thin. Septa 5/6 to 7/8 are con- ical and muscular; 8/9 to 10/11 are also muscular while 11/12 is thin. The buccal cavity opens into a muscular pharynx extending to septum 4/5. The esophagus extends from segment V to segment XV. It is un- differentiated throughout except for the unpaired, midventral sacs in X and XI, and the paired dorsolateral glands in XII. These paired glands are closely applied to 12/13 septum. The intestine begins in XVI and there are gizzards in XX-XXV. Esophageal gizzard, supraintestinal glands and typhlosole are absent. The dorsal blood vessel runs anteriorly to segment V where it breaks up into tiny vessels to supply the pharyngeal region. It gives off sub- sidiary vessels to serve the unpaired median esophageal sacs in segments X and XI. In segment XII, a large pair of vessels is given off from the dorsal vessel to serve the paired glands. This pair joins below the esophagus, separates and later joins to run posteriorly on the left side of the ventral nerve cord giving off branches to the ovo-spermathecal ap- paratus. It later becomes the subneural vessel in segment XX. Another large pair of vessels is given off in segment XIII and these run forwards to serve the paired esophageal glands. Prominent, paired septal vessels occur in 11/12 to 16/17 septa, and 2 pairs of parietal vessels are found in each intestinal segment. A supraesophageal vessel runs on the gut in segments IX to XI. Five pairs of commissural blood vessels connect the dorsal and the supraesophageal blood vessels with the ventral vessels in segments VII to XI. Each testis, in segment X and XI, is enclosed in a bean-shaped, white iridescent sac which lies near to the posterior septum of each segment. The vasa deferentia come together in segment XII and continue pos- teriorly to segment XVIII where they enter into the ental (anterior) re- gion of the euprostate. Vesiculae seminales are paired in segments XI and XII, the anterior pair extending to segment XV or XVI and posterior pair to XVI or XVII. These vesicles appear to be deeply incised. The eupros- trates are paired, whitish, muscular, cylindrical sometimes slightly curved bodies occupying segments XVII to XX on both sides of the esophagus. A long, narrow strip of muscular sheath is attached to the bodywall near VOLUME 90, NUMBER 2 261 19/20 (Fig. 2). It then runs underneath the prostatic gland and finally terminates at the male pore. This sheath contains a penial seta which is of the rolled type and which is often found protruding through the prostatic pore. Each ovary is enclosed within an ovarian vesicle from which an ovarian duct passes posteriorly into segment XIV to enter into the wider ovo- spermathecal duct. An ovisac, with a calyx-leaflike appearance at its base, leads into the fertilization chamber which opens, in turn, into the ovo- spermathecal duct through a narrow, whitish, muscular stalklike duct (Fig. 2). The spermathecal atrium is less muscular, balloon-shaped and septate. It also opens into the ovo-spermathecal duct which runs inward to open ventrally on either side of the ventral nerve cord and blood vessels through an ovo-spermathecal pore on segment XIV. Nephridia are present from segment IV. The nephridial reservoirs are comparatively larger in the clitellar region. Etymology.—This earthworm is named after Dr. (Miss) C. C. Mba, Soil Science Department, University of Nigeria, Nsukka, who sent me four preserved specimens. Acknowledgments I wish to express my gratitude to both Dr. (Miss) C. C. Mba of the University of Nigeria, Nsukka and Miss Inyang Etcheri of the University of Ife, Nigeria, for presenting some preserved earthworms from Nsukka and Calabar respectively. My thanks are also due to Joan Bacon for preparing the figures. Literature Cited Beddard, F. E. 1891. On the structure of the two new genera of earthworms be- longing to the Eudrilidae, and some remarks on Nemartodrilus. Kuart. Jour. Microscop. Sci. 32:235—-278. Clausen, M. W. 1963. Two new genera and three new species of earthworms from West Africa. Vidensk. Medd. Dansk Naturhist. Foren. 126:1—15. 1967. The African oligochaete genera Hyperiodrilus Beddard, 1891 and Legonea Clausen, 1963 (Oligochaeta). Vidensk. Medd. Dansk Naturhist. Foren. 130:179-207. Kinberg, J. G. H. 1866. Annulata nova. Fam. Lumbricina. Ofvers. K. Vetensk. Forh. Stockholm 23:98. Michaelsen, W. 1913. Oligochaten vom tropischen und siidlich-subtropischen Africa. II. Zoologica, 68:1-63. Segun, A. O. 1976. Two new genera of eudrilid earthworms from Nigeria. Proc. Biol. Soc. Wash. 88(36):383-394. In press. The genus Iridodrilus (Eudrilidae: Oligochaeta) in Nigeria. Jour. Nat. Hist. 262 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Sims, R. V. 1965. Acanthodrilidae and Eudrilidae (Oligochaeta) from Ghana. Bull. British Mus. Nat. Hist. ( Zool.) 12:283-311. 1971. Eudrilinae from Southern Nigeria and a taximetric appraisal of the family Eudrilidae (Oligochaeta). Jour. Zool. London 164:529-549. Stephenson, J. 1930. The Oligochaeta. Oxford, Clarendon Press. 978 pp. Department of Biology, University of Ife, Ile-Ife, Nigeria. PROC. BIOL. SOC. WASH. 90(2), pp. 263-273 BONADUCECYTHERIDAE, A NEW FAMILY OF CYTHERACEAN OSTRACODA, AND ITS PHYLOGENETIC SIGNIFICANCE K. G. McKenzie In general, life consistently (stubbornly, perhaps) refuses to recognize the neat compartments formulated for it by the intuitions of taxonomists whether their interests be heirarchic or phylogenetic. Only rarely do the classifications of even a decade ago satisfy the needs of researchers aware of recent advances in knowledge. In all major groups, an honest assess- ment of the situation recognizes the occurrence of taxa which are un- classifiable under the present systems. Ostracoda conform to this gen- eralization, and ostracode taxonomists are emphatic in stating the urgency of the need for a revision of the current classifications as presented by the Russian and American Treatises (Orlov, 1960; Moore, 1961). The first moves towards a new classification for Ostracoda were taken at the fifth Ostracoda Congress at Hamburg (August, 1974) with the circula- tion of a paper by Hartmann & Puri (1974), but the general ferment was already implicit in recent papers by a number of other authors (Grundel, 1967; Poulsen, 1969; Danielopol, 1972; Kozur, 1972; McKenzie, 1972; Kornicker, 1974; Kornicker & Sohn, in press). A further discussion was arranged for the sixth congress at Vienna in 1976. The Ostracode group which probably presents the greatest problem for researchers attempting to achieve a classification which satisfies both neontologists and palaeontologists, is the Podocopa. This is the largest group of living Ostracoda and it is also abundantly represented in the fossil record from the Ordovician onwards. The new family described below belongs in the Podocopa, and its description necessitates yet an- other look at the classification of that group. Phylum CRUSTACEA Linnaeus Subphylum OSTRACODA Latreille 1802 Class Podocopa Miller 1894 Order PODOCOPIDA Sylvester-Bradley 1961 Suborder PODOCOPINA Swain 1961 Superfamily CYTHERACEA Baird 1850 Family BONADUCECYTHERIDAE, new family Diagnosis.—A family of small cytheracean ostracodes characterized by the fact that in males expanded seminal vesicles are housed within the duplicature, and by the development of the unguis of the male’s first limb (PI) into a clasping structure. Except for the male PI, endopods of the 264 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON thoracic limbs have 4 segments. The carapace is extremely flattened ventrally and tapers anteriorly where it is broadly rounded. Posteriorly, it is acuminate with a distinct selvage in both valves. Anteroventrally, the valves have a pronounced gape. The line of concrescence is expressed at both the inner and outward ventral margins and is invaginated an- teriorly and posteriorly. The central muscle scar group comprises 3 ad- ductors in an oblique row and a single frontal scar. The hinge is adont. The favored environment for the taxon is interstitial. Bonaducecythere, new genus Name.—For Dr. G. Bonaduce, of Stazione Zoologica, Napoli, who col- lected the material. Type-species—Bonaducecythere hartmanni n. sp. Diagnosis——A genus of bonaducecytherid cytheraceans characterized by small size and the other features listed in the family diagnosis. Discussion.—The line of concrescence in this genus may be compared with those of the genera Microxestoleberis Miller, 1894, and Aspidoconcha de Vos, 1953, which are similarly expressed at both the inner and out- ward ventral margins. These genera, however, possess the typical arcuate eyespot and seive-type normal pore canals of xestoleberidids, characters which do not occur in the new genus. They are broader dorsally with respect to their length than is Bonaducecythere. Other genera from an interstitial habitat have been listed by Hartmann (1973) but none of them is close to Bonaducecythere either in carapace morphology or in soft anatomy. Since Aspidoconcha is a symbiont, it is worth pointing out that there are likewise few similarities between the new genus and the symbiont genera listed in McKenzie (1972b). The cytheracean families involved in these several comparisons include: Paradoxostomatidae (characterized by specialized mouthparts); Xestoleberididae (considered above); Micro- cytheridae (different muscle scars and hinge elements); Psammocytheridae (both ends rounded); Kliellidae (fused zone narrow or absent); Parvo- cytheridae (one pair less of thoracic limbs, different muscle scar pattern ); Loxoconchidae (sieve-type normal pore canals, different hinge elements); Cytheridae (sieve-type normal pore canals, few, straight radial pore canals); Krithidae (different line of concrescence as exemplified by Micro- loxoconcha Hartmann 1954)—all as diagnosed by Hartmann and Puri (1974). => Fig. 1. Bonaducecythere hartmanni A, Female carapace in dorsal view, anterior to the right, 625; B, Male left valve in lateral view, anterior to the right, «625; C, Adductors scars and frontal scar, male left valve; D, Male hemipenis; E, Fe- male PI; F, Female posterior body. VOLUME 90, NUMBER 2 265 ~L a - ° SN ° ° 266 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Bonaducecythere hartmanni, new species Figs. 1-2 Name.—For Prof. Dr. G. Hartmann, University of Hamburg, who dis- sected the female. Description—Carapace small, thin-shelled, smooth, elongate, the height little more than % the length, breadth less than half the length, right valve overlapping the left valve dorsally and ventrally, anterior broadly rounded with a flattened anterodorsal lip, posterior acuminate with 2 short postero- ventral setae in each valve, dorsum convex, venter broad and markedly flat- tened. In ventral view, the carapace is seen to gape anteriorly. Internally, the most striking feature is the very prominent selvage and the dis- tinctive line of concrescence which is deeply invaginated anteriorly and posteriorly. Radial pore canals are sometimes branched anteriorly and posteriorly, elsewhere simple and straight or curved; those at the outer ventral margin very short; normal pore canals scattered, except anteriorly where they are bunched and numerous, simple, open. Hingement adont, with accommodation for the right valve by the left. Muscle scar pattern consisting of an oblique series of 3 adductors plus a single frontal scar and (?) 2 small mandibular scars. Sexual dimorphism of the valves is not marked (Figs. 1A-C). Antennule 6-segmented; length ratios of the segments being 38:17:20:19: 13:12; first segment with a long hirsute dorsomedial bristle; second seg- ment with a shorter dorsodistal bristle; third segment with 2 dorsodistal bristles; fourth segment with 4 dorsomedial bristles and 6 dorsodistal bristles; fifth segment apparently bare; sixth segment with 2 terminal bristles (Fig. 2A). Antenna with a 3-segmented endopod; length ratios of the segments being 32:45:18; first segment with 2 short ventrodistal and 2 short dorsodistal bristles; third segment with 2 powerful terminal ungues and a weaker unguis plus a short ventrodistal bristle; exopod 3-segmented with a sensitized tip (Fig. 2C). Mandible coxale with 6 teeth decreasing in strength from front to rear, the anterior teeth with intervening “tooth- brush” bristles; epipod with 1 or 2 Strahlen; endopod 3-segmented; length ratios of the segments being 20:40:17; first segment with a clump of 4 ventrodistal bristles, at least 2 of these armed with numerous spiky hairs; second segment with 2 proximodorsal pilose bristles and 2 ventrodistal bristles; third segment with 2 terminal ungues and a terminal bristle => Fig. 2. Bonaducecythere hartmani, male. For complete chaetotaxy of limbs illustrated in this figure refer to the text. A, Antennule; B, Brush-shaped organ; C, antenna; D, Mandible; E, Maxilla epipod; F, Maxilla palp and lobes; G, PII; H, Lower lip le Pilea ete 268 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON (Fig. 2D). Maxilla comprising a 2-segmented palp and 3 lobes; the first segment of the palp carrying 4 distal bristles; the second segment with 4 terminal bristles, 2 of these unguis-like; of the lobes, the first lobe may only have 2 terminal bristles but the second and third lobes each appear to have at least 4 terminal bristles; epipod with 2 shorter Strahlen and about 10 other Strahlen (Fig. 2E, F). First thoracic limb (PI) dimorphic between male and female. Male limb comprising a protopod with a ventromedial and ventrodistal bristle; endopod 3-segmented; first segment with a ventrodistal bristle; second segment bare; third segment with a very short ventrodistal bristle and a long curved sclerotized clasping unguis rounded at the tip (Fig. 2J). Female limb (PI) with a 4-segmented endopod and a “normal” slender terminal unguis (Fig. 1E). PII and PIII non-dimorphic between males and females, each with a 4-segmented endopod and a slender terminal unguis (Figs. 2G, I). Male brush-shaped organs with 12 or more setae (Fig. 2B). Posterior of the body dimorphic, consisting in the female of paired caudal (?) processes which do not occur in the male (Fig. 1F). Genitalia of female could not be observed. Male with paired hemipenes, each characterized by a very prominent anterior processus otherwise not observed in detail (Fig. 1D). Seminal vesicles occurring as several loops apparently coiled within the posterior duplicature, a feature unique among cytheraceans. The species is blind in both sexes. Mouthparts as far as could be observed appear to be unspecialized with the mandible endopod and maxillar lobes operating as food forwarding and/or gathering struc- tures. Evidently, the anteroventral gape of the valves is a modification which facilitates ventral feeding. Dimensions.—Male: Length 0.21 mm, height 0.07 mm, breadth 0.09 mm; Female: Length 0.20 mm, height 0.06 mm, breadth 0.09 mm. Material—Two adult males and 5 adult females. Locality —St. Pauls Bay, Malta; on the south side of the Bay in front of the lido at St. Paul-il-Banar, at about 10 m out from the shore; depth 4 to 5 m; coarse and medium sized sand; the collection was made from between the bases of living Posidonia; 9 April 1974. Discussion.—In a review of interstitial ostracodes, Hartmann (1973) pointed out several adaptations which appear to be common to many taxa. These include blindness, reduction in the number of limbs and segments, reduction in reproductive parts, a distinctive shape, and alteration of some limb structures. Bonaducecythere hartmanni illustrates several of these features, in particular those related to blindness and carapace shape. The flattened carapace with a tapered anterior is well adapted to interstitial life among Posidonia. The prominent anteroventral gape of the valves is apparently unique to this taxon and seems to favor ventral feeding. The animal is evidently a detrital feeder since the mandible and mouth- VOLUME 90, NUMBER 2 269 parts are not specialized, as in Paradoxostomatidae, for symbiotic feeding on Posidonia and other marine plants. Phylogenetic Significance Before discussing the phylogenetic significance of Bonaducecythere, it is necessary to enlarge a little upon the concept of the Phylum Crustacea, since this category is a relatively new one for students of arthropods. After a lifetime of research, Manton concludes that “. . . the Arthropoda are probably polyphyletic and that arthropodisation has occurred at least three times, forming the phyla Crustacea, Chelicerata and Uniramia (Ony- chophora, Myriapoda, Hexapoda)” (Manton, 1973, p. 111). Her con- clusions have been substantiated by the results of ‘fate map’ embryological research (Anderson, 1973). Manton stresses that convergences of struc- ture are numerous between arthropodans of different taxanomic positions, and the author has also made this point (McKenzie, 1970). Once the con- cept of Crustacea as a phylum rather than as a class is considered in terms of the fossil record some previous problems disappear. Thus, no Uniramia are known with certainty before the Silurian (the marine Cambrian Burgess Shale taxon, Aysheia Walcott 1912, is an homeomorph of modern terrestrial Onychopora, not an ancestor, since it apparently lacks arth- ropodan cephalic structures and had strikingly different habits of life; e.g. it respired differently and must have fed differently). Considerations pertaining to the evolution of the predominantly terrestrial uniramians, therefore, need not cloud the perceptions of workers interested in the older predominantly aquatic crustaceans. And, since the taxonomic position of the trilobites remains anomalous, it is scarcely special pleading to suggest that more attention be paid by crustacean phylogeneticists to the Ostracoda which have the longest continuous and probably the most abundant record in the phylum. This comment is reinforced by the fact that some early fossils previously assigned to the crustacean group Phyllocarida or to riberioids have now been shown to have molluscan affinities (Runnegar and Pojeta, 1974). The distinctiveness of the Ostracoda has never been questioned and their combination into a taxon at the subphylum level follows logically from the recognition of the Crustacea as a phylum. In elevating the Podocopa to a class, the author has followed the concept of Miller in the Bay of Naples monograph (Miiller, 1894) rather than that of Sars, since for Miller Podocopa include podocopes and platycopes and this is also the author's assessment (McKenzie, 1972a). Heirarchic elevation of the Ostra- coda and Podocopa cuts out the need to create new categories, such as cohort, in order to accommodate crustacean diversity at high heirarchic levels (Dahl, 1963). Podocopida remain a group of ordinal status al- 270 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON though Sylvester-Bradley’s concept has been restricted, and, similarly, Podocopina and Cytheracea retain their levels according to the American Treatise. The significance of the new family stems from the fact that, while of predominantly cytheracean affinities, it does possess some significant fea- tures which characterize the superfamily Cypridacea. A statement such as the above must be justified in terms of taxa which could be considered to constitute typical cytheraceans and cypridaceans. Employing the chron- ological principle, since Bythocytheridae are are the longest surviving cytheraceans, comparisons will be made between Bonaducecythere and such genera as Jonesia Brady, 1866 and Bythocythere Sars, 1866. Further, since several cypridacean genera have a long fossil record in marine sedi- ments, the cypridacean comparisons will be made with such genera as Pontocypris Sars, 1866 and Macrocypris Brady, 1868 because males are known in both rather than with the about equally long-lived genus Para- cypris Sars, 1866 for which a male has yet to be described. Like other cytheraceans, Bonaducecythere has a flagelliform exopod on the antenna, unlike the small plate bearing 3 unequally long setae which occurs on Macrocypris and most cypridaceans. Further, the antennal endo- pod is not adapted for swimming by carrying long natatory setae as it is in Pontocypris and many other cypridaceans. Bonaducecythere has 3 pedi- form thoracic limbs (PI to PIII). This feature is characteristic of all cytheraceans, as opposed to the reduced PI, pediform PII and reflexed PIII which typify all cypridaceans, including Pontocypris and Macrocypris. Thirdly, Bonaducecythere either has no furca or else has a very minute furca which could not be observed following dissection. All cytheraceans, including Jonesia and Bythocythere, are similar in this respect but a prom- inent and powerful furca is typical of many cypridaceans, including Pontocypris. Whereas in cypridaceans the PI has an exopod which is usu- ally modified as a food forwarding structure and an epipod is often pres- ent, in Bonaducecythere exopod and epipod are absent and the PI is wholly pediform in the female but dimorphic and probably a clasping structure, fuctional during reproductive activity, in the male. Dimorphic thoracic limbs are not unknown in cytheraceans; witness the male PIII in Jonesia. The apparent absence of Zenkers Organs or such specialized ejaculatory apparati as occur in Pontocypris and Macrocypris is an anom- alous feature. Finally, the muscle scars and radial pore canal pattern are wholly cytheracean. On the other hand, Bonaducecythere in common with most cypridaceans has 4-segmented thoracic limb endopods (except in the male PI), a clasp- ing type unguis on the male PI and large loops of seminal vesicles coiled within the duplicature. McKenzie and Kaesler (1975) have indicated that reduction in the VOLUME 90, NUMBER 2 271 numbers of segments in individual ostracodan limbs is likely an evolu- tionary trend within the subphylum. On this assumption, Bonaducecythere is more primitive than most other cytheraceans since it has 4—rather than 3—segmented thoracic limb endopods. Another podocopine genus, Terrestricythere Schornikov, 1969, which belongs in the superfamily Ter- restricytheracea (Schornikov, 1969, n. comb. Hartmann and Puri, 1974) has a 4-segmented endopod on the walking leg (PII) and a clasping unguis on the male PI. Like Bonaducecythere, Terrestricythere is known only from modern environments. Such taxa, embodying features which may be regarded as primitive, offer a challenge to palaeontologists to find fossil representatives since they can scarcely have evolved at these high heirarchic levels within the Recent. It has been pointed out that 4-segmented PII endopods also occur in cypridaceans and it can be added that they are typical, too, of living healdiaceans and darwinulaceans. The author, on a variety of grounds, places these 3 superfamilies in the suborder Metacopina (Palaeozoic to Recent) an interpretation of relationships which have been recently criti- cized and defended (Schornikov and Gramm, 1974, Maddocks, 1973, Mc- Kenzie, in press). Then, there is the problem provided by the loops of seminal vesicles in Bonaducecythere males. Similar structures only occur in cypridaceans and the impressions of these loops on the inner valve surface are the main means by which sex dimorphism can be confirmed in fossil populations from Cenozoic assemblages. This striking similarity in part of the re- productive systems of taxa referred respectively to Podocopina and Meta- copina, throws into focus Dainelopol’s opinion that all podocopids should be considered as belonging in a single group rather than into the 2 or more subgroups into which they have been divided by other authorities (Danielopol, 1972). Summarizing, the description of Bonaducecythere hartmanni indicates no features which are unique among the Podocopida; one feature, the ex- panded seminal vesicles, which is unique among Podocopina (as the sub- order is understood by this author); and a few features which are unique for families in the Cytheracea. The totality of evidence tends to main- tain the scheme proposed by the author in 1972, except that the sub- phylum rank of Ostracoda is now specified and Podocopa and Myodocopa are elevated to class rank. It remains the author's opinion, therefore, that Ostracoda (Cambrian—Recent) in terms of living taxa may be divided into Podocopa (Ordovician—Recent) and Myodocopa (Ordovician—Recent). The class Podocopa consists of the orders Podocopida (Ordovician—Recent ) and Platycopida (Ordovician—Recent ). Podocopida comprise the suborders Podocopina (Ordovician—Recent) with the superfamilies Bairdiacea ( Ordo- vician—Recent); Cytheracea (Devonian—Recent) and Terrestricytheracea 272 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON (?-Recent); and Metacopina (Ordovician—Recent) comprising the super- families Darwinulacea (Carboniferous—Recent); Cypridacea (Devonian-— Recent); Healdiacea (Ordovician—Recent). It is necessary to add that this view does not represent a consensus: in particular, Gramm and his co-workers have criticized the placement of the living taxon Saipanetta in Healdiacea (e.g. Schornikov and Gramm, 1974). Nevertheless, the author's view is consistent with the results of a numerical cladistics study recently reported by Maddocks in which the taxa here regarded as podo- copine cluster on one branch of a Wagner tree whereas those regarded as metacopine cluster on another (Maddocks, in press). Types.—Stored at the British Museum (Natural History). Holotype ¢: BM(NH) 1976.1369. Paratypes: ?, BM(NH) 1976.1370; 13, 42, BM(NH) 1976.1371-1375. A second collection made at the type-locality is retained at Stazione Zoologica, Naples. Acknowledgments The material was collected by Dr. G. Bonaduce, of Stazione Zoologica, Napoli, Italia. The female dissection was made by Prof. Dr. G. Hartmann, University of Hamburg and the dissection of the male was completed by the author using the facilities of the British Museum (Natural History). Miss A. Scutt typed the manuscript. Literature Cited Anderson, D. T. 1973. Embryology and phylogeny of annelids and arthropods. Pergammon, London, 475 pp. Dahl, E. 1963. Main evolutionary lines among recent crustacea. Pp. 1-26 in H. B. Whittington and W. D. I. Rolfe eds. Phylogeny and Evolution of Crustacea. Spec. Publ. Mus. Comp. Zool. Harvard, xi + 192 pp. Danielopol, D. L. 1972. Supplementary data on the morphology of Neonesidea and remarks on the systematic position of the family Bairdidae (Ostracoda: Podocopida). Proc. Biol. Soc. Wash. 85:39-48. Grundel, J. 1967. Zur Grossgliederung der Ordnung Podocopida G. W. Muller, 1894 (Ostracoda). Neues Jb. Geol. Palaont. Mh., 6:321-332. Hartmann, G. 1973. Zum gegenwartigen Stand der erforschung der Ostracoden in- terstitieller systeme. Ann. Speleol. 28:417—426. , and Puri, H. S. 1974. Summary of neontological and paleontological classifi- cation of Ostracoda. Mitt Hamburgischen Zool. Mus. Inst. 70:7-73. Kornicker, L. S. 1974. Revision of the Cypridinacea of the Gulf of Naples (Ostra- coda). Smithsonian Contr. Zool. 178:64 pp. , and I. G. Sohn. 1976. Evolution of the Entomoconchacea. Pages 55-61 in G. Hartmann, ed. Proceedings of the Fifth International Symposium on Evolu- tion of Post-Paleozoic Ostracoda. Abh. Verh. Naturwiss. Ver. Hamburg, (NF) 18/19 (Suppl.):336 pp. Kozur, H. 1972. Einige Bemerkungen zur Systematik der Ostracoden und Beschrei- bung neuer Platycopida aus der Trias Ungarns. Mitt. Geol. Palaont., Innsbruck, 2: 1-27. VOLUME 90, NUMBER 2 273 Maddocks, R. F. 1973. Zenker’s organ and a new species of Saipanetta (Ostracoda). Micropaleontology 19:193—208. in press. Quest for the ancestral podocopid; numerical cladistic analysis of ostracode appendages, a preliminary report. In G. Hartmann, ed. Evolution of Post-Paleozoic Ostracoda. Manton, S. M. 1973. Arthropod phylogeny—a modern synthesis. J. Zool., Lond. 171:111—130. McKenzie, K. G. 1970. Review. Crustaceana 19:110—112. . 1972a. Contribution to the ontogeny and phylogeny of Ostracoda. Proc. IPU, XXIII int. geol. Cong., Prague, 165-188. 1972b. New data on the ostracode genera Laocoonella de Vos and Stock, Redekea de Vos, and Aspidoconcha de Vos; with a key to the family Xestole- berididae and a resume of symbiosis in Ostracoda. Beaufortia, 19:151—162. 1975. Saipanetta and the classification of podocopid Ostracoda: a reply to Schornikov and Gramm (1974). Crustaceana 29:222—224. , and Kaesler, R. L. 1975. An introduction to the numerical phylogeny and classification of paradoxostomatid Ostracoda, including a _ re-description of Machaerina tenuissima (Norman), 1869. In F. M. Swain, ed., The biology and paleobiology of Ostracoda. Moore, R. C. (Ed.) 1961. ‘Treatise on Invertebrate Paleontology. Pt. Q. Arthropoda. 3. Crustacea: Ostracoda, Geol. Soc. America, Lawrence, Kansas, xxiii + 422 pp. Miller, G. W. 1894. Ostracoden. Fauna Flora Golfes Neapel, Monogr. 21:i—viii, 1-404. Orlov, Yu. A. (Ed.) 1960. Basic Paleontology. Trilobitamorpha and Crustacea- morpha. Acad. Sci. USSR. Moscow, 515 pp. (Russian). Poulsen, E. M. 1969. Ostracoda-Myodocopa Part III A Halocypriformes—Thaumato- cypridae and Halocypridae. Dana Rep. 75:100 pp. Runnegar, B. and J. Pojeta, Jr. 1974. Molluscan phylogeny: the paleontological viewpoint. Science 186:311-317. Schornikov, E. I. 1969. A new family of Ostracoda from the supralittoral zone of the Kurile Islands. Zool. Zhurnal 68:494—498 (Russian). , and M. N. Gramm. 1974. Saipanetta McKenzie, 1968 (Ostracoda) from the northern Pacific and some problems of classification. Crustaceana 27: G2= 102" Riverina College of Advanced Education, P.O. Box 588, Wagga Wagga, N.S.W. 2650, Australia. PROC. BIOL. SOC. WASH. 90(2), pp. 274-283 OPHIOMASTIX KOEHLERI, A NEW OPHIOCOMID BRITTLESTAR (ECHINODERMATA: OPHIUROIDEA) FROM THE WESTERN INDIAN OCEAN Dennis M. Devaney In 1907, Koehler reported Ophiocoma wendti Miller and Troschel from several Indo-West Pacific localities (Fiji Islands; New Ireland; Seychelles; and Fernando Velosa, Mosambique). An illustration said to be of one of the larger specimens was given by Koehler (PL. XIII, Fig. 38) showing an example with several thickened and enlarged (claviform) upper arm spines. The distal ends of some of these spines appear to fork (furcate), a feature known to occur only among members of the genus Ophiomastix in the family Ophiocomidae. Furthermore the figure showed the upper arm plates to be mottled with dark and light color, and the arm spines banded. Attempts to examine the series of specimens reported by Koehler which are deposited in the Museum National d’Histoire Naturelle, Paris, have not been successful. Dr. G. Cherbonnier informed me (June 1970, pers. comm. ) that is was not possible to tell which of the specimens resembled the figure given by Koehler and only a single specimen said to be from Fernando Velosa was sent. That specimen with a disc diameter of 13 mm (smaller than the 20 mm maximum recorded for O. wendti by Koehler) proved to be Ophiomastix palaoensis Murakami, a species previously known only from Palau in the Western Pacific although also found in the Solomon, Marshall, and Samoan Islands (Devaney, in prep.). It now appears clear that Koehler had more than one species among those he called Ophiocoma wendti. It was pointed out in an earlier paper (Devaney, 1970:36-37) that other records of O. wendti from the Indo-West Pacific are er- roneous, based upon specimens of O. erinaceus, O. occidentalis, and O. scolopendrina, while Miller and Troschel’s O. wendti was shown to be an Atlantic (West Indian) species known more commonly in the literature as O. riisei, a junior synonym. The subsequent discovery of additional specimens from the Zanzibar re- gion of the western Indian Ocean which fitted the partial description and illustration of Ophiocoma wendti by Koehler in 1907 led to my con- clusion that a new species was represented (Devaney, 1970:37). The Zanzibar material includes a single specimen reported as O. wendti by H. L. Clark (1921) deposited in the Museum of Comparative Zoology, Har- vard (MCZ), and 2 other specimens reported and illustrated as O. wendti sensu Koehler, 1907 (pt.) by Ailsa M. Clark (in, A. M. Clark and Rowe, VOLUME 90, NUMBER 2 275 1971) in the British Museum (Natural History) (BMNH). They form the basis of the description which follows of a new species which is con- sidered a member of the genus Ophiomastix. I am most grateful to Miss Ailsa M. Clark, BMNH, for entrusting me with the specimens from that institution, to Dr. H. B. Fell for making the MCZ specimen available and to Dr. G. Cherbonnier for the specimen from the Paris museum. Ophiomastix koehleri, new species Figures 1-4 Ophiocoma wendti: Koehler, 1907 (in part) :327-328, Pl. XIII, fig 38. H. iy Clark, 1921:1129; 1938:336. Devaney, 1970:35, 37. A. M. Clark and Rowe, 1971:86-87, 91, 118-119 (note), pl. 18, fig. 5 (non O. wendti Muller and Troschel, 1842). Ophiomastix asperula (in part): H. L. Clark, 1915:294; 1921:134 (Zanzibar specimen referred to Ophiocoma wendti) [non O. asperula Liitken, 1869]. Etymology.—This species is named in honor of the French echinodermol- ogist, René Koehler, who first focused attention on it. Material examined.—Three specimens collected from the vicinity of Zanzibar Island, Indian Ocean: (a) One specimen, alcohol, disc diameter (d.d.) 23-25 mm; Zanzibar, no details; origin, Mr. M. Angel (BMNH Reg. No. 1965-6-1-451, holotype). (b) One specimen, alcohol, d.d. 22.5 mm; data _ same as for holotype (BMNH Reg. No. 1965-6-1-451, paratype). (c) One _ specimen, dry, d.d. 25 mm; Zanzibar, no data; the specimen discussed _ by H. L. Clark in 1921 as Ophiocoma wendti (MCZ No. 1795, paratype). Diagnosis.—Disc granulated, without spinules. Presence of some modi- fied claviform-furcate upper arm spines. Two tentacle scales regularly only on distal segments. Upper arm plates variegated dark and light, arm _ spines banded. Description of holotype (BMNH Reg. No. 1965-6-1-451).—d.d. 23-25 mm; arms about 5 times d.d., all broken before tips. Upper surface of disc covered with rounded granules. A random assortment of 25 granules from the disc average 136 wm in diameter (range = 101-177) and 163 wm in height (range = 139-202). In no case is difference between height and diameter of individual granules greater than 50 pm; in all cases but one (where diameter and height are the same) granule height is greater than its diameter. Granules relatively uniform in distribution, generally not touching each other; radial shields covered by granules; granules continue as broad V-shaped area interradially, becoming sparse near oral shields; 3 or 4 slightly conical granules at distal edges of oral shield; a series of OF WASHINGTON VOLUME 90, NUMBER 2 277 Fig. 2. Ophiomastix koehleri, BMNH paratype. < Fig. 1. Ophiomastix koehleri, BMNH holotype: A, Upper surface; B, Lower surface. 278 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 3. Ophiomastix koehleri, MCZ paratype: A, Upper surface; B, Lower surface. similar or even more tapering granules along interradial edge of genital opening (these may be in double row part way from oral shield); no evidence of disc spinules.1 Oral shield noticeably longer than broad, (1.3:1.0) with greatest breadth distal to middle of shield, only gradually narrowing inward to form straight broad front; madreporite shield wider, with median distal convexity. Adoral Shields widely separated within by oral shield; inner radial angle concave VOLUME 90, NUMBER 2 279 or straight, with buccal tentacle scale only touching near distal part of the angle; outer radial angle touching ventral shield. Three oral papillae on each side: outer papilla widest with inner side concave; its proximal edge touching adjacent papilla; inner papilla slightly larger than middle papilla, but both inner and middle papillae smaller and not as wide as outer. Buccal tentacle scale contiguous at distal edge with adoral and ventral shields, projecting into mouth slit and partly covered by outer oral papilla. One or 2 subdental papillae at apex of and between oral (jaw) plates. Thirteen to 15 dental papillae to each jaw, in 3 rows with 3 or 4 ir- regularly placed papillae deeper. Teeth with wide, blunt hyaline tips. Lower arm plates near disc to middle of arm broader than long (1.8: 1.0) with broadly rounded distal border and tapering distal angles, some- times slightly concave at mid-distal border; % of total breadth of arm plate separating lateral arm plates at distal border; arm plates becoming more elongate on distal segments. Upper arm plates with irregular lateral borders where upper arm spines alternate (4 and 3) being truncated where 4th (upper) spine encroaches; upper arm plates with strongly tapering lateral margins and slightly con- vex median proximal border on segments with 3 spines on each side. The number of arm spines irregularly alternating beyond disc margin; the sequence for number of arm spines beginning with the first arm seg- ment (beneath the disc) is as follows: Part beneath disc Part beyond disc Segment cre 4, Soko. te 6 Oo TO" Wi to 3” 14 1S 16 Number COMMON Ronin mitaweAys G4 fy 4. ecSre S40 She 44 end 4 of CMON On NOD oat pes Geka A ee 2 3 A Arm Spines Ba B Oe Rory ait As” 4" "4. ~— "4 4 3 Seo Oto ae Siro) roa woune oni 4a Alisotny4 9 Sup Boyt Sra 3 CML OU oe Seek Ou kk ee tee Aa ot. A BA SOM Oe TO. woe a a ee Ae a S| SS CeeOP mor roe oe ane caren Te eae Sara Ss OM toinsruron ire ries! midye iA Shucde 4 3 { Sun Shy hOue [oan Suen The upper (4th) arm spines on segments beyond disc differ in shape and size from lower spines in same row or from upper (3rd) spine on adjacent or Opposite segments (Fig. 4A, B) by being much larger, from 4 to 5 seg- 280 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 4. Ophiomastix koehleri, holotype, arm spines: A, 4th (upper) modified spine (left) and 3rd spine (right) from same row on segment 14; B, 3rd (upper) spine from segment 15; C, 4th (upper) spine from segment 24; D, 4th (upper) spine from segment 45; E, 3rd (upper) spine from segment 88 with bifurcate tip (left), and 3rd (upper) unmodified spine from segment 89 (right). A, B, E, 3.5; GD <4. ments long, usually somewhat swollen in central part then tapering before expanding into a flattened often forked (bi- or tri-lobed) tip (Fig. 4C); from approximately segment 40, there are fewer segments with 4 spines and the 3rd (upper) spines may be enlarged (Fig. 4D) and forked, fol- lowed by segments with 3 or 2 spines in a row in which the upper spine is unmodified and located in a lower position on the lateral plate than the modified spine; a modified upper (3rd) spine from segment 88 is compared with an upper (3rd) spine from segment 89 in Figure 4E. Two tentacle scales on each side of arm segments regularly only on first 8 to 12 segments; increasingly fewer segments with 2 scales and more often only 1 scale on distal segments; where 2 scales occur, outer is larger, but same length as inner scale; where a single scale occurs it is larger than either of the 2 scales and is rounded with a slight taper toward tip, but becomes more tapered on distal segments of arm. Pigmentation (in alcohol): Basic colors dark blackish brown and white with following pattern: disc above and below uniformly dark brown includ- ing granules, except that there is some brown and white color in interradial areas beginning at oral shields; granules upon white areas in interradial region and those along edge of genital apertures also white. Upper arm plates mottled irregularly dark brown and white, rarely is a plate a single color, frequently blotches or bands of dark color predominate, leaving only distal border white. Similar colors on upper side of lateral arm plates. Lower arm plates more regular in pattern with proximal part brown and distal margin white; proximal part of lateral arm plates brown continuing into the brown on lower arm plates. Tentacle scales brown and white, brown in 2 or 3 irregular bands across scales with edges white. Arm VOLUME 90, NUMBER 2 281 spines, except for enlarged upper ones, distinctly banded brown and white with brown rings slightly wider than white rings; many of the spines with tips white; upper modified spines more uniform in color, light brown but with some annulation toward the tips. Mouth region showing consider- able brown color especially oral shields with only their outer margins white; a part of each adoral plate brown, as well as parts of oral papillae and inner edge of oral plates. (A part of the disc and arms of the holotype is shown as Plate 18, figure 5 by A. M. Clark and Rowe, 1971.) Comparison with Other Specimens and Variations A. Disc cover—BMNH paratype (d.d. 22.5 mm) similar to holotype in distribution of granules, but with only 1 or 2 granules at base of oral shield. A random assortment of 25 granules was examined: the granules averaged 124 wm (89-139) in breadth and 117 pm in height (101-139). In contrast to the holotype the granules were generally slightly broader than high or equal in height and diameter, the ratio never exceeding 1.0: 1.4. The MCZ paratype (d.d. 25 mm) is similar to the holotype. B. Tentacle scales —BMNH paratype with 2 scales regularly on each side to segments 12, 11, 8, 9, 13; with 1 scale becoming more frequent on distal segments. MCZ paratype with 2 scales regularly on each side to segment 15, 11, irregularly to segment 25 and on more distal segments 2 scales widely spaced, otherwise 1 scale. C. Arm spine sequence—BMNH Paratype: Segment oe iA 25) (16 Vas (Or OE lOO Lye 1B) TAS NS Number Sie Ole BOL Fon CO AeA AAG AGS, eA eo tena of Ore COO Oe Owe ee ee Ae Aa we a Arm Spines SIO LOM to) Mutter. # ALe TAM AAS si fila Ye) UA Ou BON AO | hon ety Ae ee) why Al A a a ee Seer Ou OM Pom OMe Aas eae eA Ae RA eet en Ae Sek A Se Sdn toom. Sema ies Bh Se Ws eC ae Sema! in? Sa Siew Or MAO wee ae ae Or GOs doo TOL a 4 Brecon ka Ohh aA MCZ Paratype CMe rour Cote ore Ge oO. ao a 282 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Segment LQ) 8 46.455. 6) “hen tS (3:9) olOn Tel Oasis ee re Number of SYat) ee eae area kee ee tale | a7! Arm Si meOMe Oo) Leone RE ETA aeois Ly Spines WwW WwW Www GC) eH) C9 CH C5 CD Www Www Www WwW Www wor PP PR PE aN Comparing these arm spine sequences with that of the holotype one notes that all but one side of the first 4 arm segments carry 3 arm spines and the 5th segment carries 3 and/or 4. The presence of 5 arm spines on any side of an arm segment is rare but may occur between segments 7 and 10. The irregular alternation of 3 and 4 arm spines usually is evident from segment 10. The MCZ specimen shows modified enlarged upper arm spines from segments 9 to 51 on one ray examined. These modified spines occur on opposite sides of segments 9 and 10, both sides of segments 12, 14, and 17, then on one side of segments 20, 21, 24, and 25. Farther out on arm they occur on every 2 to 4 segments. Some spines are distinctly forked at tip while others are only flattened at tip and somewhat swollen in middle of spine. Some spines reach up to 6 segments in length. Two arm spines were noted on a few segments beyond segment 20, being most evident past segment 40. Segments beyond 40 also showed the 3rd (upper) spine in a row enlarged and elongate; from this part of the arm to the end, the presence of 4 arm spines is infrequent and 3 spines are more common. In the same row on a segment side, the lowest spine is usually as long as, or somewhat longer than, the next spine and is slightly curved. D. Pigmentation—BMNH paratype with aboral surface of disc brown where granules are present; underlying scales lighter; granular area over radial shields lighter brown in a narrow band. Lower arm plates, beyond segment 10, having dark brown color in 2 lateral bands with center light except for small dark spot near distal border; sometimes central part of plate also dark. Tentacle scales with several dark spots. Mouth shields with several brown blotches; adoral shields with medial brown spot. Inter- radial part of disc, from oral shield out to granular area, with light and dark scales, some of the granules white. Some upper arm plates have brown color in blotches, but otherwise as described for holotype, with majority having the distal edge white. Arm spines not completely banded, rather VOLUME 90, NUMBER 2 283 with alternating bands of brown and white on upper and lower sides but interrupted with light line along lateral edges. The MCZ paratype is quite similar to the holotype in color. Relationships The new species, Ophiomastix koehleri, strongly links Ophiomastix with the Scolopendrina group of Ophiocoma. Both Koehler (1907) and H. L. Clark (1921) rightly considered that this form as Ophiocoma wendti combined characters of both genera. The low, rounded, non-contiguous granules, absence of disc spinules, and alternation of arm spines present in Ophiomastix koehleri are char- acteristic of the Scolopendrina group of Ophiocoma (Devaney, 1970). However, the latter is also a feature of Ophiomastix, while the presence of enlarged upper arm spines, with the tips forked is known only among species of Ophiomastix and is the criterion which leads me to place this new species in this genus. Of those species of Ophiomastix possessing disc granules, only O. asperula Liitken has these more abundant than spinules. However, the granules of O. asperula are conical and not bluntly rounded as in O. koehleri. Literature Cited Clark, A. M., and F. W. E. Rowe. 1971. Monograph of shailow-water Indo-West Pacific echinoderms. British Mus. (Nat. Hist.) London. 283 pp., 31 pls., 100 text-figs. Clark, H. L. 1915. Catalogue of recent ophiurans based on the collection of the Museum of Comparative Zoology. Mem. Mus. Comp. Zool. Harvard 25(4): 165-376, 20 pls. 1921. The echinoderm fauna of Torres Strait: Its composition and _ its origin. Carnegie Inst. Washington Publ. No. 214:1-223, 38 pls. Devaney, D. M. 1970. Studies on ophiocomid brittlestars. I. A new genus (Clark- coma) of Ophiocominae with a reevaluation of the genus Ophiocoma. Smith- sonian Contrib. Zool. 51:1-41, 50 figs. Koehler, R. 1907. Revision de la collection des Ophiures du Muséum d Histoire naturelle de Paris. Bull. Scient. France et Belgique 41:79-351, pls. 10-14. Division of Invertebrate Zoology, Bernice P. Bishop Museum, Honolulu, Hawaii 96819. Footnote * Granules are herein defined as elements whose height is 2 times or less than their diameter or greatest breadth; spinules as elements whose height exceeds twice the maximum diameter or greatest breadth. PROC. BIOL. SOC. WASH. 90(2), pp. 284-290 NOTES ON THE BIOLOGY OF THE POTONIINE SHRIMP LIPKEBE HOLTHUIS! CHACE, WITH A DESCRIPIMON OF THE MALE J. Kevin Shaw, Richard W. Heard, Jr., and Thomas S. Hopkins Chace (1969) first described the pontoniine shrimp, Lipkebe holthuisi, from two female specimens collected from the Gulf of Mexico, off the coast of Florida at a depth of 119 m. A second occurrence of this shrimp was reported by Bruce (1976a) from off Brazil; only a single specimen, an ovigerous female, was noted. To our knowledge no male has been dis- covered. This paper reports on a series of collections of L. holthuisi from the eastern Gulf of Mexico. Included is a description of the male from three specimens, and the commensal relationship of L. holthuisi with the crinoid Comactinia meridionalis meridionalis (Agassiz) from 150 m depth. We wish to thank Drs. Fenner A. Chace, Jr., and Thomas E. Bowman, Department of Invertebrate Zoology (USNM) and Austin B. Williams, National Marine Fisheries Service Systematics Laboratory for critically reading the manuscript and making available type-material. We express our gratitude to Ms. Linda Lutz and Mr. Tom Walker for preparation of the figures. We wish to thank Dr. David L. Meyer, University of Cincin- nati for the crinoid identifications. This work was supported by a sub- contract of the Bureau of Land Management Contract No. 8550-CT5-30 to Dr. Thomas S. Hopkins. Lipkebe holthuisi Chace, 1969 Figures 1-2 Lipkebe holthuisi Chace, 1969:263-266, figs. 8-9; 1972:25.—Bruce, 1976a: 310-312, figs. 1-2. Material examined.—Oregon Station 1024, 1 ovig. 2 (holotype, USNM 97433), 25°13’N 83°55’W, 119 m, 19 April 1954. MAFLA Station 15-I-B, 1 ovig. 2, 26°25’N 83°49’W, 150 m, 28 July 1975. MAFLA Station 33-I-B, 7 females (5 ovig., 1 with bopyrid isopod), 2 males (3.3 mm, 2.8 mm USNM 168535), 26°25’N 83°50’W, 150 m, 28 Feb. 1976. MAFLA Station 46-310201, 2 females (1 ovig. 3.6 mm, 2.9 mm USNM 168534), 1¢, 26°24'30’N 83°49/30”W, 150 m, 15 July 1976. Morphology.—The ovigerous female specimens agree with the descrip- tion of Chace (1969) and no additions are included. However, the male specimens of Lipkebe holthuisi differ from the females in the size and shape of the carapace and second pereiopods (Fig. 1). As in the ovigerous female, the rostrum extends to the distal end of the antennular 285 NUMBER 2 VOLUME 90, Fence ”/ Sa ~ Fig. 1. Lipkebe holthuisi: a, 286 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON peduncle and is laterally compressed in the distal half. When viewed laterally, the carapace is more in line with the rostrum of the male than in the ovigerous female, where it appears distended. The carapace is re- duced in length relative to the abdomen and much more depressed than in the ovigerous female; it is compressed laterally. The lateral antennular flagellum of the larger male appears slightly longer than that of the fe- male; it is nearly twice the length of the internal antennular flagellum. The second pereiopods of the male are, as in the female, very unequal and dissimilar. However, the major cheliped of the male is less robust, the palm being compressed laterally. It over-reaches the antennal scale by the chela and only part of the carpus. The fingers are less than half as long as the palm in the smaller specimens, increasing to slightly longer than half the palm in the largest male. The dactyl is similar to that of the female, having a large basal tooth closing into a depression within the fixed finger. The merus, carpus, and propodus contain the minute tubercles depicted by Chace (1969: fig. 9b, 3). The minor cheliped is slightly shorter than that of the female, extending beyond the antennal scale by less than the full length of the chela. As in the female, the fingers are 0.7 as long as the palm and unarmed. Bruce (1976a) noted that the small spinules on the ventral border of the dactyls of the ambulatory pereiopods (Chace, 1969: fig. 9g, i, k) are not discernible. These are distinctly present in all specimens from the Gulf of Mexico. The specimen from Brazil also seems to have many more long, coarsely plumose setae in groups on the distal ends of the propodus of these limbs than do the specimens from off the coast of Florida. The immature female resemble the male and is distinguishable only by its secondary sexual characters. These characters are illustrated for both the male and female in fig. 2. The margin of the endopod of the first male pleopod is entire, not bilobate. Up to 3 small setal spines (asetes) project medially from the margin. The appendix masculina on the second pleopod of the largest male has a setal combination of 3, 3, 6; apical, subapical, lateral (Fig. 2c). The apical setae originate from a straight tip and extend by % their length beyond the small, rounded ap- pendix interna. The setation of the first and second pleopods of the males is summarized in Table 1. These follow the characters evaluated by Fleming (1969) for male external genitalia of certain species of Palae- monetes. As noted in Table 1, addition of setae occurs with an increase in body size of the male. The setae present on the appendix interna on the second pleopod of females (Fig. 2e, f) range from none in the small, im- mature females to 6 in the largest ovigerous female. As in the male, the numbers of setae increase as the body size of the female increases (Table 2) VOLUME 90, NUMBER 2 287 Fig. 2. Lipkebe holthwisi: a, Male, first pleopod; b, Male, second pleopod: ec, Male, appendix masculina; d, Female, first pleopod; e, Female, second pleopod; f, Female, appendix interna. a, b, d, e, Scale = 1.0 mm; c, f, scale = 0.1 mm. 288 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Table 1. Male characters of the first and second pleopods of Lipkebe holthuisi. Number of setae eae Eee second pleopod, appendix masculina (mm ) endopod lateral subapical apical Si) 3 6 3 3 2.8 2 6 1(2)* 2 2.8 lt 5) 1 2 * (2) from right pleopod. Remarks.—The carapace lengths of the males range from 2.8 to 3.3 mm. The females range 2.9 to 4.6 mm. One female bears an undescribed bopyrid isopod of the genus Bopyrina Kossman, 1881, within its right branchial chamber. This species is being described by one of the authors (RWH). In comparing the shrimp with the non-parasitized females, no alterations of the secondary sex characters were found; however, this is the only non- ovigerous female over 3.0 mm in length. The color of live Lipkebe holthuisi is a deep crimson hue, blending naturally with its commensal host, Comactinia meridionalis meriodionals. Some shrimp were preserved within the arms of the crinoid, which was collected in large numbers. A single living shrimp was observed with its host for a short time in an aquarium. It moved about only on the oral plate and not on the arms of the crinoid. Distribution—Lipkebe holthuisi occurs in the western North Atlantic Table 2. Female characters of the second pleopods of Lipkebe holthuisi. Carapace second pleopod, appendix interna length (mm) (number of setae) on) 4.6 4.4*+ 4.2 3.9 3.9 3.8 3.6 3.4 2G) Dae) SON WwW Kk WY W OL WU * Non-ovigerous. + With branchial bopyrid isopod. VOLUME 90, NUMBER 2 289 from the southeastern Gulf of Mexico and the western South Atlantic off the coast of Brazil; 64-150 m depth. Off the coast of Florida it is as- sociated with the crinoid Comactinia meridionalis meridionalis. Discussion—The genus Lipkebe appears to be related to the genera Pontonides Borradaile, Neopontonides Holthuis, Veleronia Holthuis, and Pseudocoutierea Holthuis, which are associated with gorgonians and anti- patharians, and the probably commensal genus Coutierea Nobili, whose host is not known (Bruce, 1976a; Chace, 1972). From the similarity in form of the dactylus of ambulatory pereiopods in the aforementioned genera, Bruce (loc. cit.) suggested a similar type of host for L. holthuisi. The association of L. holthuisi with a crinoid may extend the possibilities for the commensal host of Coutierea. The accounts of caridean shrimp commensal on crinoid hosts are well documented. Potts (1915) included the behavioral activity and color vari- ations of the Pontoniinae and Alpheidae commensal on their crinoid hosts. More recently, Bruce (1965; 1971) provided invaluable information on the crinoid associates of the Indo-Pacific species of Periclimenes and other pontoniine-crinoid associations. Fishelson (1974) observed the cryptic behavior and coloration of three pontoniine shrimp on crinoids from the Red Sea and noted their mode of feeding. In his review of the commensalism of coral reef Caridea, Bruce (1976b) listed 5 genera of the Pontoniinae and the genus Synalpheus as commensals occurring on crinoid hosts. Additional observations on the alpheid associations with the crinoid Comatularum group from the Indo-Pacific were discussed by Banner and Banner (1975), who included notes on color variations, behavioral activity, and host specificity. Recent records of pontoniine-crinoid associa- tions from the Western Atlantic are found in the descriptions of Peri- climenes bowmani, P. crinoidalis, and P. meyeri by Chace (1972). Of interest is the specificity of Lipkebe holthuisi to the crinoid Comac- tinia m. meridionalis. A smaller, yellow crinoid, Leptonemaster venustus Clarke, co-occurs with C. m. meridionalis, but no shrimp were collected from it. All specimens of L. holthuisi were colored to blend with their host, a deep crimson. There was no variability in color of the shrimp to suggest a different or additional host, as indicated by Potts (1915). Literature Cited Banner, D. M., and A. H. Banner. 1975. The alpheid shrimp of Australia. Part 2. The genus Synalpheus. Rec. Australian Mus. 29(12):267-389. Bruce, A. J. 1965. Notes on some Indo-Pacific Pontoniinae. X. periclimenes cristi- manus sp. nov., a new pontoniid shrimp from Singapore. Ann. Mag. Nat. Hist. 13(8) :487—493. —. 1971. Periclimenes attenuatus sp. nov. (Crustacea, Decapoda, Natantia, Pontoniinae), a new commensal shrimp from Duke of York Islands. Pacific Sci. 25:533-544. 290 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 1976a. A further occurrence of Lipkebe holthuisi Chace (Decapoda, Natantia, Pontoniinae). Crustaceana 30(3):310-312. 1976b. Coral reef Caridea and “Commensalism.” Micronesia 12(1):83-98. Chace, F. A., Jr. 1969. A new genus and five new species of shrimps (Decapoda, Palaemonidae, Pontoniinae) from the Western Atlantic. Crustaceana 16(3): 251-272. —. 1972. The shrimps of the Smithsonian-Bredin Caribbean Expeditions with a summary of the West Indian shallow-water species (Crustacea, Decapoda, Natantia). Smithsonian Contrib. Zool. 98:1-178, figs. 1-61. Fishelson, L. 1974. Ecology of the northern Red Sea crinoids and their epi- and endozoic fauna. Mar. Biol. 26:183-192. Fleming, L. E. 1969. Use of male external genitalic details as taxonomic characters in some species of Palaemonetes (Decapoda, Palaemonidae). Proc. Biol. Soc. Wash. 82:443-452. Potts, F. A. 1915. The fauna associated with the crinoids of a tropical coral reef: with a special reference to its color variation. Carmegie Inst. Wash. Pap. Dept. Mar. Biol. III:73—97. Dauphin Island Sea Lab, P.O. Box 386, Dauphin Island, Alabama 36528. Footnote * Contribution Number 105 of the Marine Environmental Sciences Consortium, Dauphin Island, Alabama 36528. PROC. BIOL. SOC. WASH. 90(2), pp. 291-301 THE IDENTITY OF LEWIS’ MARMOT, ARCTOMYS LEWISII Robert S. Hoffmann To which mammal the name Arctomys lewisii Audubon and Bachman, 1848, should be attached has remained uncertain for many years. The genus Arctomys Schreber is now known as Marmota Blumenbach, but at the time Arctomys lewisii was named, the genus also included species presently recognized as belonging to the genera Ammospermophilus, Cyn- omys, Spermophilus and others. The original description of A. lewisii was based on a specimen from a locality ambiguously given as “Columbia,” and the describers assumed that the specimen had come from the vicinity of the Columbia River, in the northwestern United States. However, the description and the type-specimen do not conform to any species known to occur in the Columbia River region. It is the purpose of this paper to identify Arctomys lewisii, and to discuss its confused taxonomic history. I believe that the holotype did not originate in North America at all, but instead is a specimen of Marmota baibacina, the Central Asian montane marmot, for reasons presented below. The Original Description In volume 3 of “The Viviparous Quadrupeds of North America,” the Reverend Doctor John Bachman (Audubon and Bachman, 1846-53) de- scribed a new species of marmot, Arctomys lewisii Audubon and Bach- man, 1848, based on specimen number 461 in the Zoological Society of London collection, and now a skin and skull in the British Museum ( Nat- ural History), numbered 55.12.24.144 in their catalog. Bachman’s written description was preceded by a color plate (CVII) based on an original study executed by John Woodhouse Audubon, the younger son of Bach- man’s co-author John James Audubon, from the type-specimen in London, and issued in volume 3 of the Imperial Folio (Audubon and Bachman, 1845-48). Subsequently, plates and text were published together in an octavo edition retitled “The Quadrupeds of North America,” volume 3 appearing in 1854 (Audubon and Bachman, 1849-54). The name Arctomys lewisii is usually dated from 1853 ( Baird, 1857; Allen, 1898) or 1854 ( Hollis- ter, 1916; Hall and Kelson, 1959). However, the name first appeared on the plate published in 1848, and according to the International Code of Zoolog- ical Nomenclature, a name published before 1931 becomes available if accompanied by a description, definition, or indication. An indication in- cludes publication of the new name “. . . in connection with an illustra- tion; ...’ Hence, the name takes date and authorship from 1848 ( Audubon and Bachman, 1845-48). 292 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON As the specimen was attributed to “Columbia” both on the skin label and in the Zoological Society catalog, Audubon and Bachman interpreted this as “. . . the shores of the Columbia River ... ,” naming it in honor of Captain Meriwether Lewis, first American explorer of that region. Subsequent Attributions Although Bachman unequivocally stated that the specimen possessed “the characteristics of the true Marmots,” S. F. Baird (1857) called attention to the fact that Bachman described the thumb of A. lewisii as long, with a long nail. Both he, and subsequently J. A. Allen (1877 and 1898) sug- gested, on the basis of the supposedly well-developed, clawed hallux of the type-specimen, that it was not a Marmota at all, but rather a prairie dog (Cynomys), and J. A. Allen (1877:903) first proposed that the name Arctomys lewisii be applied to Cynomys columbianus (= C. gunnisoni), and later (1898:456), to C. leucurus, over which it had priority. How- ever, neither Baird nor Allen examined the type-specimen. It fell to Ned Hollister, while revising the genus Cynomys, to query Oldfield Thomas concerning the type of Arctomys lewisii. Thomas replied, according to Hollister (1916:26) that the specimen in question was “...a species of Marmota, and not of Cynomys as supposed by Dr. Allen.” (The original correspondence is, unfortunately, lost. ) There the matter rested, and Arctomys lewisii has, when attributed at all, generally been regarded as a junior synonym of Marmota flaviventris (Audubon and Bachman). This is plausible in view of the reddish- orange ventral coloration visible in J. W. Audubon’s illustration, and the specimen’s supposed provenance; the question of the long, clawed hallux described by Bachman has not, it will be noted, been answered yet. The Type-Specimen Some time ago, during the course of an extensive systematic survey of the genus Marmota, I had an opportunity to examine the type-specimen of Arctomys lewisii, and to compare it with specimens of other taxa of Marmota in the British Museum collections, as well as with my own rec- ords on the marmots of a number of major collections in the Soviet Union, western Europe, and the United States. I unhesitatingly concurred with Oldfield Thomas that the specimen was assignable to Marmota, and not Cynomys. However, the skin had all toes severed at the base on all four feet, and was thus without claws. I will return to this discrepancy between the published description by Audubon and Bachman, and the present condition of the type-speci- men. Even though it lacked toes, I recognized the specimen as Marmota baibacina Kashchenko, 1899, a species that inhabits the mountains and VOLUME 90, NUMBER 2 293 foothills of Central Asia, from the central Tyan Shans to the western Mongolian Altai (Ognev, 1947:279). The skin and skull are those of a young animal, probably about one year old and killed in its second spring of life, judging from the pelage and the fact that the third upper permanent premolar is not yet fully erupted. In pelage color and pattern, A. lewisii resembles M. baibacina rather than the New World marmots. Marmota baibacina, and its close relatives in the Old World (M. bobac, M. siberica, M. himalayana, M. camtschatica, M. menzbieri) differ from all the New World Marmota, except arctic M. broweri in having dorsal guard hairs uniformly dark both at the base and tip, with a lighter middle band. Moreover, M. baibacina is charac- terized by its orangish ventral coloration, and brown head, lacking a pro- nounced dark cap, with the brown color extending down the sides of the head and neck to meet the orange throat. Audubon and Bachman’s description of the color of the type-specimen agrees in most respects with that of M. baibacina, for they note that “, . . the longer [dorsal] hairs, at their extremities [are] blackish brown .. . feet and belly, light salmon-red; tail, from the root for half its length, reddish-brown, the other half to the tip soiled white ... .” J. W. Audubon’s illustration carefully reflects all of these characteristics of M. baibacina, and considering that he painted it from a museum skin, compares favor- ably with the illustrations of M. baibacina in Bobrinskii, et al. (1965) and Flint, et al. (1965). In contrast, Audubon and Bachman (1841:29) described the dorsal fur of M. flaviventris as “. . . on each hair a con- siderable space is occupied by dirty yellowish-white, which is gradually shaded towards the apex through brown into black tips of hairs yellowish- white...” I also compared the skull of the type with skulls of approximately the same age of M. baibacina and M. flaviventris available in the British Museum. In qualitative characters such as shape of the nasal bones and post-orbital region, relative interorbital width, and curvature of the profile of the skull, Arctomys lewisii resembles M. baibacina and not M. flavi- ventris. This is also true of several measurements (Table 1). Thus, in most features, the type-specimen and its description agree with the characteristics of Marmota baibacina. One discrepancy, however, is the terminal “soiled white” tail tip, conspicuous in Audubon’s illustration, which led Allen (1898) to suggest that Arctomys lewisii might be a white- tailed prairie dog (Cynomys leucurus). My examination of the type re- vealed that the distal half of the tail retains the previous year's old, un- molted hairs. These are extremely faded, and contrast strongly with the rest of the fresh spring pelage. This retention of faded, unreplaced fur on the rump and tail is a frequent occurrence in the molt pattern of Marmota (Kapitonov, 1964). 294 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Table 1. Comparison of selected cranial dimensions for “Arctomys lewisii” with Marmota f. flaviventris and M. baibacina centralis. Dimension, mm M. flaviventris “A. lewisii” M. baibacina Length, mand. tooth row 18.4 22.4 21.9 Length diast. Upper 16.0 19.4 19.6 Lower 10.0 13.6 WAS Width, p4 4.8 oul 3.4 Length, p4 5.4 4.3 4.2 Width, m3 5.5 5.5 6.0 Length, m3 6.7 Tol 7.9 The Question of the Thumb The second major discrepancy between the description of Arctomys lewisii and M. baibacina, or indeed, any Marmota, is that the “. . . thumbs, instead of being remarkably short and equipped with blunt nails, have long nails nearly the length of those on the other toes” (Audubon and Bachman, 1854:31). Moreover, toe coloration is described, and the mea- surement of hind foot length (heel to middle claw) is given. Yet the digits are now wanting on all four feet. One possibility is that Bachman accurately described the condition of the digits of the type-specimen at that time, but that the toes have been removed since then, a highly unusual procedure. This possibility leaves the long, clawed hallux unexplained. The other possibility is that the type- skin was originally without toes, and that the details in the description were added for some reason—in this case the long thumb with its claw never existed. A case can be made for this second possibility, if the his- tory and context of the description of Arctomys lewisii are considered. Bachman’s “Discovery” of Arctomys lewisii Bachman was in London only once, after visiting J. J. Audubon and his family in Edinburgh, in the summer of 1838. He worked in the collections of the British Museum and the Zoological Society of London for several weeks before going on to Germany, and returned home in January, 1839 (C. L. Bachman, 1888:175; Ford, 1964:354). J. J. Audubon and Bachman published, in 1841, “Descriptions of new species of quadrupeds inhabiting North America,” based in part on specimens examined in the Zoological Society of London, presumably in 1838. Among these is Arctomys (= Marmota) flaviventer Audubon and Bachman (1841:29), based on a “*.. . VOLUME 90, NUMBER 2 295 specimen in the [Zool. Soc. Lond.] collection brought by the late David Douglass [sic] . . . from the mountains between Texas and California, and is marked in their printed catalogue of 1839, “Arctomys flaviventer. No. 459, Bachman’s Mss.” (op. cit.:30). The printed catalog referred to is a supplement to the catalog published by Waterhouse (1838). In the 1838 catalog, the following North American marmots are listed: “459. Quebec Marmot ... From North America. Arctomys Empetra. Schreb. Presented by Dr. Richardson. 459a. Ditto ditto ... Habitat North America. 460. Whistler . . . Habitat Rocky Mountains. Arctomys? pruinosus. Rich. Died in the Menagerie. Presented by B. King, Esq. 461. Short-tailed Marmot . . . Habitat Columbia. Arctomys brachyurus? Harlan.” Specimen number 459a of the 1838 catalog was designated the type of Arctomys flaviventer by Audubon and Bachman (1841); they also men- tion, in “The Quadrupeds of North America,” having examined specimen no. 459 (vol. 1, pg. 24, 1849), and specimen no. 460 (vol. 3, pg. 19, 1854). It would be reasonable to assume that Bachman also examined specimen no. 461 during that same period in 1838. This was not the case. On 31 October 1846, Victor G. Audubon, John James Audubon’s elder son, wrote to Bachman, requesting the scientific names to accompany the plates of the sea otter, musk ox, “whistler,” “Columbia pouched rat,” “hare from Texas,” and “short-tailed marmot” (letter 262, by permission, Houghton Library, Harvard Univ.). Bachman replied in a letter to J. J. Audubon, dated 5 November 1846, giving the names of the first five, and continuing: “. . . 6. Short tailed marmot. Now friend—here is fun. By some unaccountable means I never saw the specimen in England. I am deeply mortified about it. It has never been described Lewis & Clarke mention it. No specimens were brought. Harlan named it Arctomys? brachyurus—short tailed—named it so after Lewis & Clarkes description. Now we must name it, but alas I dont know whether it is an Arctomys or Spermophylus—I am quite in a quandary. Perhaps we had better wait till we hear from Waterhouse through John [Woodhouse Audubon, J. J. Audubon’s younger son]. The specific name—brachyurus of Harlan agree- able to our rule we must not take, besides it is an improper one as its tail is larger than many others. Cant you or Victor do this. Just send me an outline of the drawings dabbling a little of the colours to give me an idea of it. I strongly suspect it must be a spermophile. Write to John at once 296 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON & let him ask Waterhouse to examine it—but let him not hint that it is not described as those Zool. boys will name it an hour after. O that I had wings for an hour & telegraphic speed to carry me to London—but here Iam & cant move... .” (Copy of letter in Charleston Museum, Charleston, South Carolina; a variant version of this letter appears in C. L. Bachman, 1888:224.) Apparently Bachman’s request was not complied with promptly enough to suit him. In a letter dated 14 December 1846, to Victor Audubon, he complained “. . . You promised to send me a lithograph of the short tailed marmot till then I am in the dark—I dont even know whether it is a Spermophil—John & Waterhouse ought to be consulted Now for your plates . . . 107. Short tailed marmot No name till I see Te pa All of these are clearly references to “Arctomys brachyurus,” and the MS plate number (107) corresponds to the number of the plate of Arctomys lewisii that was finally published. On 27 December 1846, Bachman wrote again to Victor “. . . Above all say to John that we wish the names from Waterhouse of the species he is fig- uring. I have great doubts about some of them. I wish the short tailed Marmot to be examined in regard to the Genus. I wish John was as willing to write as he is to paint but John wont do it, what shall I say more... .” (letter, Charleston Mus.). I have not yet been able to determine whether Victor or John W. Audu- bon wrote to Bachman concerning A. lewisii, but he did receive the promised lithograph. A letter from Victor, dated 24 March 1847, referring to another request from Bachman concerning a deer, assures him that “When we have a proof [plate] I will send it to you so that you may see it as you did the short-tailed marmot.” (letter 264, Houghton Library, Harvard University). I find no further reference to Arctomys lewisii in the subsequent Audubon-Bachman correspondence I have seen; it appears that Bachman’s questions concerning the “short-talied marmot” were re- solved between 27 Dec. 1846 and 24 Mar. 1847. It may be that the litho- graph proof was the only descriptive material that Bachman received. Imperial Folio plate 107 reveals a clawed hallux on the specimen of A. lewisii much more clearly than the smaller, redrawn octavo edition. Unfortunately, the location of J. W. Audubon’s original study is unknown (Ford, 1951:215). Bachman was initially inclined to the position that “Arctomys brachy- urus’ was a spermophile, but was assured by Victor and John that it was a marmot, Arctomys. He later decided that the name given the specimen in the catalog “Arctomys? brachyurus Harlan,” actually applied *. . . to some species of spermophile—probably Spermophilus townsendii ... (Audubon and Bachman, 1849-1854, vol. 3, pg. 34), and that Zoological VOLUME 90, NUMBER 2 297 Society specimen no. 461 thus represented an undescribed species of marmot. Origin of the Type-Specimen of Arctomys lewisii The remaining substantive question is: How did a specimen of M. bai- bacina happen to arrive at the Zoological Society of London under such circumstances that it was labeled “Habitat Columbia,” and that J. W. Audubon was told, apparently, that the specimen “. . . was sent to the Zoological Society by the British fur-traders who are in the habit of annually carrying their peltry down the Columbia River to the Pacific” (Audubon and Bachman, 1849-1854). The Zoological Society collection was started by the “Zoological Club” in 1823 (Scherren, no date; Thomas, 1906). The specimen was not listed in a catalog of the collection published in 1829 (Anon.), but was nine years later (Waterhouse, 1838); the period during which the specimen might have arrived in London is thus defined. One possibility is that the specimen actually was obtained by British fur-traders. Pelts from the North West Company, which later merged with the Hudson Bay Company, were shipped from Astoria on the Columbia River, across the Pacific to London, on ships which often called at Chinese ports (Davidson, 1918:164). The trade in furs between Russia and China was of long standing, and “Furs constituted an extraordinarily large share of the goods exported by the Russians to China... .” (Fisher, 1943:224). Hence, it is possible that the skin of a M. baibacina taken in the Tyan Shan or Altai might make its way from a Russian trading town to a Chinese port, there to be picked up by a British trader homeward bound to London. The principal reason to doubt such a chain of events is the condition of the type-specimen of A. lewisii. Although the digits are severed, most of the skin of legs and feet is present, and the cuts in the skin are not the sort that would have been made by a hunter pelting a marmot to produce a skin for market (see Louashkin, 1937, Pl. XCVII). Finally, one would not expect a commercial skin to remain associated with its cor- responding skull throughout the sort of journey hypothesized above. Since the condition of the specimen suggests that it was obtained with its scientific significance in mind, with what scientific collections might a specimen of M. baibacina become associated during the time period in question (1823-1838)? One possibility would be the several collections made by David Douglas in western North America between 1824 and 1833 (McKelvey, 1955). Douglas, it will be remembered, was the collector who obtained the specimen of Marmota flaviventris, no. 459 in the Zoolog- ical Society collection, that was first described by Audubon and Bachman. 298 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Douglas in 1831 visited Fort Ross, in what is now northern California, and later corresponded with Baron Wrangel, governor of Russian America (Mor- wood, 1973). It is possible that a Russian marmot came into his possession in this connection, and was later sent to London with his other specimens. Another possibility involves the ship Blossom, commanded by Captain F. W. Beechey; the naturalist aboard was George T. Lay, and the surgeon, Alexander Collie. The Blossom left England on 19 May 1825, and eventually reached Petropaulski, Kamchatka, on 27 June 1826. There it met with Baron Wrangel’s ship Modeste and stayed five days. The Blossom visited Petropaulski a second time on 3 July 1927, staying this time 15 days; it arrived home on 12 October 1828 (Beechey, 1831). During the course of their two visits to the Siberian port, Lay and Collie might have obtained, by gift or exchange, a Russian marmot specimen, which later arrived in London with the other scientific collections obtained during the course of the voyage (Richardson, 1839). A third possibility is that it was part of a collection of Russian mammals obtained by either the Zoological Society or the British Museum from the collector I. G. W. Brandt of Hamburg, sometime prior to 1843. Brandt sent to the British Museum a number of mammal specimens, including a “Tamias striatus” [= T. sibiricus|] and “Spermophilus altaicus” [= S. undu- latus| from the Altai Mountains, and a “Marmota bobac” from Siberia (Gray, 1843). Moreover, there is considerable similarity in manner of preparation between the “M. bobac’ obtained from Brandt, and the type- specimen of Arctomys lewisii, which differentiate both specimens from many others prepared during that period. For example, in addition to a mid-ventral cut, both specimens have cuts around the bases of three legs; wire rather than wood was used to support the tail; the stuffing material— straw with bits of cotton—is similar; and the stitch pattern used to sew the cuts is alike in both. Also, both specimens are shaped similarly, with a rounded, slightly elevated head distinctly set off from the body by a con- striction in the neck. The plantar surfaces of the forefeet face up, and those of the hind feet, down. Thus, it seems possible that the specimen of “M. bobac’ received from Brandt, and the type-specimen of Arctomys lewisii, may have been prepared by the same person. I have not found any record of purchase of specimens from Brandt by the Zoological Society. However, the first year that Brandt sold specimens to the British Museum was 1840. Since Gray only began to register speci- mens in the British Museum catalog in 1837 (Thomas, 1906), Brandt is not likely to have had a market for his specimens there prior to that date, and it is possible that he sold some to the Zoological Society, which was still actively building its collections in the period 1829-1838 (Scherren, no date). VOLUME 90, NUMBER 2 299 Taxonomic Implications It is unlikely that a satisfactory determination of the way in which the type-specimen of Arctomys lewisii reached the Zoological Society of London will ever be made. By whatever means this skin and skull of Marmota baibacina reached London, its designation as the type of A. lewisii by Audubon and Bachman must be reckoned with. J. F. Brandt (1843) first applied the name Arctomys baibacina to a specimen of marmot from the Altai Mountains of south-central Siberia. Unfortunately, he did not describe the specimen. This lapsus was corrected in 1899, when Kashchenko recognized that M. baibacina was a nomen nudum, and renamed and described the species. In the meantime, Audubon and Bachman had named and figured (1848), and subsequently described (1853) a specimen of M. baibacina although they attributed it to the wrong continent. The name M. lewisii thus has priority over M. baibacina. However, due to the many uncertainties surrounding it, and the likeli- hood that the specimen will never be accurately ascribed, nomenclatural stability is not served by replacing the name baibacina with lewisii. More- over, lewisii has been virtually unused since it was proposed, except for its misapplication by J. A. Allen (1898), following Baird (1857) as a synonym of Cynomys leucurus. Under the terms of Article 23b of the International Code of Zoological Nomenclature, “A name that has remained unused as a senior synonym in the primary zoological literature for more than fifty years is to be considered a forgotten name (nomen oblitum).” Accordingly, I have re- ferred the name Arctomys lewisit Audubon and Bachman, 1848 to the International Commission on Zoological Nomenclature, for inclusion on the Official List of Rejected and Invalid Specific Names in Zoology. Acknowledgments Parts of this study were begun while I was a U.S. National Academy of Science exchange fellow with the Soviet Academy of Sciences. It was continued at the universities of Montana and Kansas, and completed as a visiting fellow at the National Museum of Natural History, Washington, D.C. The following persons have contributed material help to the study at one stage or another: R. G. Dennis (Houghton Lib., Harvard U.), L. I. Galkina (Biol. Inst., Novosibirsk), A. Gardner (Nat. Mus. Nat. Hist., Washington, D.C.), I. M. Gromov (Zool. Inst., Leningrad), J. E. Hill (British Museum [Nat. Hist.], London), V. Kapitonov (Inst. Zool., Alma- Ata), B. M. Petrov (Biol. Inst., Belgrade; formerly Mus. Nature, Tashkent), A. E. Sanders (Charleston Museum, S.C.), and P. L. Wright (U. Montana). B. H. Blake reexamined the holotype of A. lewisii and British Museum 300 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON (Natural History) records several times, to answer my specific questions as they arose, and criticized the manuscript, as did H. Setzer (Nat. Mus. Nat. Hist., Washington, D.C.), and G. Corbet (B.M.[N.H.], London). Without the help of these persons and others, this study could not have been completed. Literature Cited Allen, J. A. 1877. Monographs of North American Rodentia. No. XI.—Sciuridae. pp. 631-940 in Rept. U.S. Geol. Surv. Territories, F. V. Hayden, ed. Wash- meton. Ch x —— x= 1091 pp: 1898. Nomenclatural notes on certain North American Mammals. Bull. Amer. Mus. Nat. Hist. 10:449-461. Anonymous. 1829. Catalogue of the animals preserved in the museum of Zoolog- ical Society, Sept. 1829. Richard Taylor, London. 40 pp. Audubon, J. J., and J. Bachman. 1841. Descriptions of new species of quadrupeds inhabiting North America. Jour. Acad. Nat. Sci. Phila. Ser. 1, 8:1-43. 1845-1848. The viviparous quadrupeds of North America. Vol. 1, pl. 1-50; Vol. 2, pl. 51-100; Vol. 3, pl. 101-150. J. J. Audubon, New York. 1846-1853. The viviparous quadrupeds of North America. Vol. 1, xvi + 390 pp:; Vol. 2, 336 pp.:; Vol. 3, vi + 257 pp. J. J. and V.)@ sAudubonteNew, York. 1849-1854. The quadrupeds of North America. Vol. 1, viii + 383 pp.; Vol. 2, 334 pp.; Vol. 3, v + 348 pp. V. G. Audubon, New York. Bachman, C. L. 1888. John Bachman. Walker, Evans and Cogswell Co., Charleston, S.C. xi + 436 pp. Baird, S. F. 1857. General report upon the zoology of the several Pacific railroad routes. Pacific R.R. Rept. 8. Pt. 1. Mammals. Washington, 757 pp., 60 pl. Beechey, F. W. 1831. Narrative of a voyage to the Pacific . . . in the years 1825, 1826, 1827, 1828. 2 Vol. Henry Colburm and Richard Bentley, London. Bobrinskii, N. A., B. A. Kuznetsov, and A. P. Kuzyakin. 1965. Opredelitel’ mlekopitay- ushchikh SSSR. “Prosveshchenie,” Moscow. 382 pp., 40 pl., 111 maps. Brandt, J. F. 1843. Observations sur les differentes especes des Sousliks de Russie. Bull. Classe Phycico-mathem. Acad. Sci. St. Petersbourg, 3( 23-24) :357-382. Davidson, G. C. 1918. The North West Company, Univ. Calif. Publ. Hist. 7. xi + 349 pp. Fisher, R. H. 1943. The Russian fur trade 1550-1700. Univ. Calif. Publ. Hist. ol:xni + 1-276. Flint, V. E., Yu. D. Chugunov, and V. M. Smirin. 1965. Mlekopitayushchie SSSR. “Mysl’,” Moscow. 437 pp., 40 pl. Ford, A. 1951. Audubon’s animals. The quadrupeds of North America. The Studio Publ., New York. 222 pp. 1964. John James Audubon. Univ. Okla. Press, Norman. Gray, J. E. 1843. List of the specimens of Mammalia in the collection of the British Museum. Brit. Mus., London. xxviii + 216 pp. Hall, E. R., and K. R. Kelson. 1959. The mammals of North America. 2 Vols. Ronald Press, New York. xxx + 546 + 79; viii + 547-1083 + 79 pp. Hollister, N. 1916. A systematic account of the prairie-dogs. N. Amer. Fauna, No. 40. 37 pp., 7 pls. Kapitonov, V. I. 1964. Lin’ka surkov. AN Kazakh. SSR, Trudy Inst. Zool. 23:169- 190. VOLUME 90, NUMBER 2 301 Kashchenko, N. F. 1899. Rezul’taty Altaiskoi ekspedetsii. Tomsk, ii + 158 pp., A pl. Loukashkin, A. S. 1937. The tarbagan or the Transbaikalian marmot and its economic value. Comptes Rendus, XII Cong. Intern. Zool. Lisbon, 1935. pp. 2233-2293, pl. XCV-XCVII. McKelvey, S. D. 1955. Botanical exploration of the Trans-Mississippi west. Arnold Arboretum, Harvard Univ., Jamaica Plain, Mass. xi + 1144 pp. Morwood, W. 1973. Traveler in a vanished landscape. C. N. Potter, New York. x + 244 pp. Ognev, S. I. 1947. Zveri SSSR i prilezhashchikh stran. Vol. 5. Gryzuny. Akad. Nauk, Moscow-Leningrad. 809 pp. Richardson, J. 1839. Mammalia, pp. 1-14 in The Zoology of Captain Beechey’s voyage ... in the years 1825, 26, 27, and 28. H. G. Bohn, London. Scherren, H. No date. The Zoological-Society of London .... Cassell & Co., London. Thomas, O. 1906. The history of the collections contained in the Natural History Department of the British Museum. 1. Mammals. Publ. Brit. Mus. 2:3—66. Waterhouse, G. R. 1838. Catalogue of the Mammalia preserved in the Museum of Zoological Society of London. 2nd ed. R. and J. E. Taylor, London. 68 pp. 1839. Supplement 13 pp. Museum of Natural History and Department of Systematics and Ecology, The University of Kansas, Lawrence, Kansas 66045. PROC. BIOL. SOC. WASH. 90(2), pp. 302-325 THE MILLIPED GENUS CROATANIA (POLYDESMIDA: XYSTODESMIDAE) Rowland M. Shelley The xystodesmid milliped fauna of the lowland regions of the Caro- linas is sparse in comparison to that of the southern Appalachian Moun- tains. Apheloria tigana Chamberlin is dominant in the fall zone region of North Carolina (Shelley 1977), and Sigmoria latior (Brolemann) (three subspecies plus intergrades) ranges through piedmont North Caro- lina to the southern Coastal Plain of South Carolina (Shelley 1976). Cleptoria macra Chamberlin is known from the counties of Green- ville and Newberry, S.C. (Hoffman 1967), and Pachydesmus crassicutis incursus Chamberlin occurs from the inner Coastal Plain and Piedmont of South Carolina to the Kings Mountain region (Cleveland and Gaston counties) of North Carolina (Shelley and Filka 1977). Other lowland xystodesmids include Nannaria conservata Chamberlin, N. rutherford- ensis Shelley, and Pleuroloma sp. in North Carolina (Shelley 1975, 1977), and Stelgipus agrestis Loomis in South Carolina (Chamberlin and Hoff- man 1958). In August 1975 I discovered several xystodesmids in a moist seepage area in the Kings Mountain, N.C., region that represented an undiagnosed generic form. A year later the genus was found to be common in a va- riety of piedmont South Carolina biotopes ranging south to the Savannah River. Available museum specimens confirm its occurrence on islands of the extreme southern coast of the state. Four distinct species are repre- sented, with no evidence of intergradation or development of geographic races. Accordingly, I propose the name Croatania for this complex, and present descriptive, ecological, distributional, and phylogenetic informa- tion. Materials and Methods Over 170 specimens have been examined during this study, and except for five in the personal collection of Richard L. Hoffman (RLH), all were collected by the author or an assistant and deposited in the invertebrate collection of the North Carolina State Museum (NCSM), the invertebrate catalog numbers of which are indicated in parentheses. Color and habitat observations were recorded in the field at the time of collection. Illustrations were prepared with the aid of a binocular stereomicroscope fitted with a grid reticle, and measurements were made with vernier-scale calipers of specimens lying flat, straight, and compressed. VOLUME 90, NUMBER 2 303 The format for this publication is similar to the xystodesmid revisions of Hoffman, as exemplified by his work on Cleptoria (1967). Thus, a dis- cussion of taxonomic characters precedes the descriptive sections, there being no prior literature to review. Discussions of ecology, distribution, and phylogenetic relationships, both generic and specific, conclude the paper. Illustrations of diagnostic features and a taxonomic key are pro- vided to aid in determinations and establish the identities of the species. Taxonomic Characters Color.—The paranotal color of many xystodesmids is highly variable and a poor taxonomic character, but in Croatania, it is remarkably constant for each species. All specimens of C. saluda, for example, have red paranota, whereas both C. catawba and C. simplex display a yellowish color. There may or may not be a concolorous stripe along the anterior edge of the col- lum, and some individuals of C. saluda have narrow paranotal markings cov- ering little more than the peritremata. Two specimens of C. catawba have an orangish hue to the paranota, but this could not possibly be con- fused with the red of C. saluda. Thus, color affords a general basis for field identifications of the three Piedmont species; the color of C. yemassee, in the Coastal Plain, is unknown. Sterna.—The postgonopodal sterna of Croatania are a modification of the bilobed variation described by Hoffman (1965). The true bilobed condition, producing acute subcoxal spines, is most evident on segments S-ll of C. catawba, and the sterna become progressively flatter pos- teriorly. In the other species, the bilobed condition is only noticeable on segment eight. The pregonopodal sterna exhibit several modifications. On the fourth segment of both sexes there is a large, ventrally directed, bilobed process (Figs. 11-15), which is most highly developed in males of C. catawba and C. saluda. Here it is longer than the widths of adjacent coxae, bent anteriad distally, and divided into two lobes near the apices. In males of C. yemassee and C. simplex the structure is straight, and in the latter it is shorter than the width of the adjacent coxae and divided near mid- length. In all females the process is much shorter than the adjacent coxae and almost completely separated into two paramedian lobes (Fig. 1») Segments five and six of males also possess sternal modifications. On segment five, between the anterior (4th) legs, is a pair of digitiform lobes which are subsimilar in all four species. Between the posterior (Sth) legs there are elevated flattened areas that also do not vary appreciably between species. The sternum of segment six is recessed between the seventh legs to accommodate the distal curvature of the gonopodal telopo- dite. This depression is relatively shallow in C. catawba, C. saluda, and C. 304 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON simplex, but much deeper in C. yemassee due to the higher arc of its telopodite. Gonopods.—As with most xystodesmids, the principal diagnostic struc- tures at both the generic and species levels are the male gonopods. Neither coloration nor sternal configuration can alone distinguish Croatania from related genera. Two features of the gonopods are especially important—a large pre- femoral process, and an elaborate, irregularly notched expansion of the proximomedial edge of the telopodite. The latter, which imparts a bizarre, spiny appearance to the gonopods, is unique to Croatania and presumably a specialization. It varies in length and degree of jaggedness and is re- duced and relatively inconspicuous in C. yemassee. The prefemoral pro- cesses of C. catawhba, C. saluda, and C. yemassee (Figs. 1, 3, 6) are enor- mous, extending to or beyond the level of the tip of the telopodite, and set them apart from sympatric xystodemids. The fourth species, C. sim- plex, differs in having a reduced prefemoral process that terminates well below the level of the tip of the telopodite (Fig. 4). Additional features of the gonopods that are diagnostically important in- clude the degree of arch of the telopodite, the sizes of the medial and lateral subapical lobes, and the length of the solenomerite relative to the lobes. In C. catawba and C. saluda the gonopods are so similar that there is little doubt about their cogeneric status. The other two species, C. yemassee and C. simplex, differ sufficiently, however, that one could justifiably question their being congeneric with each other as well as with the more typical species. The former, in addition to a greatly reduced proximomedial expansion of the telopodite, has a much higher distal arch and more reduced subapical lobes than either C. catawba or C. saluda. The prefemoral process, however, is equivalent in size to those of the latter species, and this, plus the presence of subapical lobes and a proximo- medial expansion, even though reduced, justify inclusion in the same genus. The fourth species, C. simplex, is even more dissimilar, and I originally considered erecting a monotypic genus to accommodate it. As indicated above, this species is unique in having a short prefemoral process, but it is also distinguished by an elongate, acuminate solenomerite. The thin, reduced subapical lobes of the telopodite are even more inconspicuous than those of C. yemassee. Their presence, however, plus the well de- veloped, notched expansion and the occasional appearance of bifurcation in the prefemoral process, which resembles the condition in C. saluda, are enough, in my opinion, to place the form in the same genus. This is the conservative approach, especially since C. simplex is known from only one locality, and additional material might provide a clearer indication of the relationship. Further discussion of relationships in Croatania is presented in the concluding section. VOLUME 90, NUMBER 2 305 Cyphopods.—The operculum and valves of the female cyphopods of Croatania are small and surrounded on three sides, ventrally, anteriorly, and caudally, by the receptacle. The membrane to which the cyphopod is attached is greatly enlarged and folded, particularly on the medial side, where it protrudes through the aperture (Fig. 16). This bulging medial enlargement appears to be a fundamental part of the female anatomy, although its function is unknown. It is most highly developed in C. catawba, but present in females of all four species. In some individuals it con- tains darkened inclusions. This structure has never before been reported in the Xystodesmidae and is considered a clear indication of generic dis- tinction for this complex. It is also further justification of congeneric status for C. simplex. Croatania, new genus Type-species.—Croatania catawba, new species. Description—A genus of moderate to large xystodesmids with the fol- lowing characteristics: Body composed of head and 20 segments in both sexes; relatively large and compact in form, W/L ratio varying from 21 to 26 percent. Head of normal appearance, smooth, polished; epicranial suture distinct, not bifid; interantennal isthmus relatively broad; genae not or moderately im- pressed; facial setae reduced, epicranial and interantennal absent, with or without frontal setae. Antennae extending caudad to posterior portion of third metatergite; with four antennal cones on ultimate antennomere; no other sensory structures. Terga polished, smooth medially, becoming coriaceous near paranota; latter broad, depressed, continuing slope of dorsum; peritremata flat on anteriormost segments, becoming more distinct and elevated in midbody region; Ozopores opening dorsolaterad. Prozonites generally smaller than metazonites, meeting at distinct sutural line, becoming most sharply de- fined at level of stricture. Caudal segments normal for family. Sterna variable, either flattened on postgonopodal segments or depressed medially and produced into prominent subcoxal lobes, lobed condition most prominent on eighth segment; sternum of fourth segment produced into large medial process; of fifth segment, produced into two prominent digitiform lobes between fourth legs; recessed on sixth segment between seventh legs. Gonopodal aperture subelliptical, narrowing slightly dorsolaterad. Gono- pods large, projecting forward beyond edge of aperture into sternal de- pression between seventh pair of legs, usually not overlapping. Coxa large, without apophysis, connected by membrane only, no sternal remnant. Prefemur small, with large prefemoral process arising on dorsal side. Telop- 306 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON odite with irregularly notched expansion along proximomedial edge, curved anteriad, apex recurved with subapical medial and lateral lobes; solenomerite curved proximad, length variable. Cyphopods with convoluted enlargement of membrane protruding through medial portion of aperture; receptacle visible in lateral half of aperture, extending around valves; valves directed dorsad, subequal in size; operculum minute, concealed under free end of valves. Range.—Coastal Plain and Piedmont physiographic provinces of North and South Carolina, from Lincoln Co., in south-central North Carolina, to the coastal islands of southern South Carolina (Fig. 17). Species.—Four. Additional species may be encountered in the Piedmont province of Georgia. Key to Species of Croatania (based on adult males) 1. Prefemoral process of male gonopod large, extending to or be- yond level of tip of telopodite 2 — Prefemoral process reduced, terminating well below level of tip of telopodite; Chester Co., S.C. simplex, new species 2. Prefemoral process apically bifurcate; Newberry to McCormick counties, S.C. saluda, new species — brefemoral process simple, not bifurcate 3 3. Prefemoral process subglobose basally; proximomedial edge of telopodite with conspicuous, irregularly notched expansion; arc of telopodite relatively flat and broad; distal extremity of telopodite subequal to outermost point in curvature of prefemoral process; sub- apical lobes of telopodite large, conspicuous; Lincoln Co., N.C. to Union Co., S.C. catawba, new species — Prefemoral process of approximately uniform width except near tip; expansion of proxiomedial edge of telopodite inconspicuous, with only small denticulations; arc of telopodite relatively short and high; distal extremity of telopodite extending beyond outermost point in curvature of prefemoral process; subapical lobes reduced, indistinct; coastal islands and outer Coastal Plain of Beaufort and Jasper counties, S.C. yemasseé, new species Croatania catawba, new species Figs. 1-2, 7, 11-12, 16 Type-specimens.—Male holotype (NCSM A540) and eight male and five female paratypes collected by R. M. Shelley and J. C. Clamp, 16 August 1975, from Cleveland Co., N.C., 5.8 mi S Kings Mountain VOLUME 90, NUMBER 2 307 Figs. 1-6. Left male gonopods of Croatania spp., medial views. 1-2, C. catawha; 1, Holotype; 2, Prefemoral process of specimen from Woods Ferry Recreation Center, Chester Co., South Carolina. 3, C. saluda, holotype. 4—5, C. simlex; 4, Holotype; 5, Bifurcate prefemoral process of paratype. 6, C. yemassee. 308 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON (town), along county road 2245, 0.1 mi N junction county road 2288. Male and female paratypes deposited in Florida State Collection of Arthro- pods, Zoologisch Museum Amsterdam, and personal collection of Richard L. Hoffman. Diagnosis.—Paranota yellow, with or without yellow stripe along an- terior edge of collum; sternal processes between third pair of legs of males longer than width of adjacent coxae, diverging near apices, bent anteriad distally; postgonopodal sterna distinctly bilobed on segments 8-11; gono- pods with following characteristics: prefemoral process subglobose basally, bent sharply ventrad at % length; expansion of proximomedial edge of telopodite relatively short, irregularly notched; arc of telopodite relatively flat and broad; subapical lobes of telopodite large, distinct; solenomerite subequal in length to lateral lobe; distal extremity of telopodite subequal to outermost point in curvature of prefemoral process. Holotype —Length 50.6 mm, greatest width 11.4 mm; W/L ratio 22.5%. Paranota depressed, creating high depth/width ratio of 61.4% at mid- body. Body essentially parallel sided in midbody region, tapering at both ends as indicated by following segmental widths: Collum 8.0 mm 14th 11.0 Mids A932 15th 10.5 3rd 9.9 16th 9.9 Ath-6th 10.7 17th 8.9 8th-12th 11.4 18th 6.4 Color in life-——Paranota bright lemon yellow, metaterga black; broad yellow stripe along anterior edge of collum, connecting with paranotal spots. Epicranium, genae, and frons black, epicranial suture and clypeus lighter; antennae brown, ultimate segment darkest. Venter and legs light brown, claws and distal ends of tarsi darker. Head capsule smooth, polished; width across genal apices 5.3 mm; inter- antennal isthmus broad (1.7 mm), smooth; epicranial suture thin but dis- tinct, terminating at level of antennae. Antennae moderately long and slender, extending to middle of paranota of third segment, becoming pro- gressively more hirsute distally; first antennomere subglobose, 2-6 clavate, 7 short and truncate. Genae not margined laterally, trace of medial im- pression, ends broadly rounded and extending slightly beyond adjacent margins of cranium. Facial setae as follows: epicranial and interantennal absent; frontal 1-1; genal 3-3; clypeal about 8-8, irregularly spaced; labral about 12-12, varying in length. Terga polished, coriaceous. Collum large, broad, extending slightly be- yond ends of following tergite on each side. Paranota depressed, con- VOLUME 90, NUMBER 2 309 tinuing slope of middorsum, corners rounded on segments 2-4, becoming slightly more distinct on 5-7 and progressively more pointed posteriorly due to thickening of peritremata. Peritremata virtually flat through segment 6, only slightly elevated above paranotal surface; higher and more distinct on segments 7-8. Ozopores small and indistinct, situated near middle of peritremata, opening dorsolaterad. Sides of metazonites irregular, with several shallow, curved impres- sions. Strictures broad, distinct. Sternum between third pair of legs pro- duced into large process (Fig. 11), divided apically, bent anteriad distally, extending well beyond margins of adjacent coxae; sternum of segment 5 produced into two small digitiform lobes between fourth legs and into two broad elevated areas between fifth pair; sternum of segment 6 re- cessed slightly between seventh legs. Postgonopodal sterna a modifica- tion of bilobed variation discussed by Hoffman (1965); elevated behind stricture, interrupted by transverse groove behind first pair of legs, with distinct medial vertical groove on segments 8-10, becoming shallower, rounded, and less distinct on segment 11; acute subcoxal spines sub- tending both pairs of legs on segments 8-11, becoming indistinct at an- terior legs on segment 12 and gradually more plate-like behind posterior legs of each segment. Hypoproct rounded; paraprocts with edges strongly thickened. Coxae generally without spines, low blunt tubercles present on legs 15-25; prefemoral spines distinct on legs 6-30, indistinct on first five legs; tarsal claws bisinuately curved. Gonopodal aperture subelliptical, narrowing slightly dorsolaterad, about 4.5 mm wide and 2.4 mm long at midpoint; margins raised, slightly thickened along caudal edge. Gonopods (Figs. 1, 7) large, extending for- ward just beyond anterior edge of aperture, not overlapping. Coxa massive. Prefemur small; enormous prefemoral process arising on dorsal side of prefemur, subglobose basally, widest just proximal to midlength and blackened along inner edge, bent ventrad at about % length and tapered to acuminate tip, not bifurcate; tip directed toward midpoint of curve of telopodite, extending beyond apex of telopodite. Telopodite with proxi- momedial edge expanded into deeply notched plate, basalmost projec- tion enlarged into wedgelike structure; telopodite curved broadly anteriad distal to expansion, distalmost point at approximately same level as outer- most point in curvature of prefemoral process; with subapical lobes on each side, lateral thick and pointed, larger than medial, latter thin and rounded; solenomerite short and blunt, bent caudad, length subequal to lateral lobe, longer than medial lobe; raised ridge with tuberclelike projection about * length along inner curvature of telopodite marking course of prostatic groove; groove running mostly along lateral edge of ridge, crossing over from medial side near base of pretemoral process. Male paratypes—The W/L ratios of the eight male paratypes range 310 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON from 22.0%—22.8%. All specimens display the stripe along the anterior edge of the collum and prominent sternal processes between the third pair of legs. The postgonopodal sterna vary little from the condition in the holotype. On the gonopods, the configuration of the proximomedial ex- pansion varies, but the basalmost projection is always the largest. The tuberclelike projection of the ridge along the inner face of the telopodite also varies, being low, rounded, and virtually absent on some individ- uals, and sharply pointed and spiniform on others. The prefemoral pro- cesses are remarkably similar; the relative thickness of the basal sub- globose portion varies slightly, and some are bent ventrad more sharply than others. Female paratypes——The five female paratypes agree closely with the males in body proportions, except that they have a higher depth/ width ratio, the average being 71.5%. Three individuals lack the yellow stripe along the anterior edge of the collum. The sternal process between the third pair of legs (Fig. 12) is much smaller than those of males, being completely divided and shorter than the width of the adjacent coxae. Only a pair of subcoxal spines is present between the fourth legs; the sterna on segments 5-7 are as on segments 8-10 of the holotype. On the cyphopods (Fig. 16), the medial enlargement of the membrane is very large and conspicuous, and contains darkened inclusions. The receptacle appears striate and extends almost completely around the valves. The latter are small, subequal, and closely adhered together; the operculum is minute and concealed under the free end of the valves. Variation.—The paranotal color of this species is bright lemon yellow on all but two individuals, a male and female from Union Co., S.C., which displays an orange tint. The stripe along the anterior edge of the collum may be present or absent, but when present is usually broad. As noted earlier, all nine males and two of the five females of the type-series possess the stripe, whereas it is absent from two females. In a large, ran- domly collected sample of 35 adults (276 ¢, 822) from Chester Co., S.C., the stripe is absent from 22 males and all eight females. Thus, only 18.5% of the males and 14.3% of the total sample possess this trait, which is a rough indication of its frequency in the general population at this site. In addition to the stripe along the anterior edge of the collum, a few individuals also possess thin, faint stripes along the caudal edges of the metaterga of segments 1-4. These stripes are only present on individuals which also display the one along the anterior edge of the collum. The body proportions of the holotype and male paratypes were re- corded earlier. A table summarizing the dimensions of the sample from Chester Co. is given below. The remainder of the non-sexual charac- ters are as described for the holotype. VOLUME 90, NUMBER 2 311 W (mm) L (mm) W/L (%) N Range 9 “Range Range Males 27 99-122 111 39.9-51.6 46.0 21.1-24.8 22.5 Females 8 AO) = 1OIe Galatea 46.3-51.3 48.1 99.4-95,.1 23.1 The gonopods are remarkably constant, the most notable variation being the degree of jaggedness of the proximomedial expansion and the prominence of the projections of the mid-telopodite ridge. The males from Chester Co. differ, however, in having much thinner prefemoral pro- cesses which are not subglobose basally (Fig. 2). This variation does not have any geographical significance and seems to be characteristic of this particular population. Ecology—The type specimens were collected from a seepage area along a small feeder stream to Kings Creek on a rocky hillside in a climax deciduous forest (elevation approximately 1,000 ft). Though a hot, dry August day, the site was comparatively cool and moist, and deep piles of leaves had accumulated beside some of the larger boulders. Several individuals were discovered under these leaf piles, while others were found under small logs or moving across the ground. The species ap- peared to be abundant, and more material could have been secured. It was not encountered at several other localities in the Kings Mountain area, even though the habitats are similar to the type locality. This site was revisited in late November, 1975, and April, 1976, but no specimens were found. The South Carolina specimens were also taken during hot, dry August weather and were especially abundant in the predominantly hardwood forest at the Chester Co. site, along the Broad River. Distribution.—Croatania catawba is known from the south-central Pied- mont of North Carolina and the north-central Piedmont of South Carolina (Fig. 17). Specimens have been examined as follows: North Carolina: Lincoln Co., 7.3 mi W Lincolnton, 1¢, 25 October 1952, L. Hubricht (RLH). Cleveland Co., 1.2 mi SE Kings Mountain (town), along I-85 at KOA campsite, 56, 12, 6 juvs., 10 July 1976, M. Filka and W. W. Thomson (A1049); and 5.8 mi S Kings Mountain (town), along co. rd. 2245, 0.1 mi N jet. co. rd. 2288, 96, 52, R. M. Shelley and J. C. Clamp (A540) TYPE-LOCALITY. Gaston Co., 4.8 mi SW Gastonia, along co. rd. 1131, 2.1 mi NW jet. co. rd. 1133, 12, M. Filka and W. W. Thomson (A1340). South Carolina: York Co., Kings Mountain State Park, 3¢, 32, 20 August 1976, R. M. Shelley (A1416). Chester Co., 13.6 mi W Chester, Woods Ferry Recreation Center, campsite on Broad River in Sumter National Forest, along U.S. Forest Service rd. 574, 3.0 mi W jet. S.C. Hwy. 312 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 49, 278, 82, 2 juvs., 4 August 1976, R. M. Shelley (A1414). Union Co., 12.3 mi SE Union, along S.C. Hwy. 121-72 at crossing Cane Cr., 46, 72, R. M. Shelley (A1415). Croatania saluda, new species RSS y 4 Oils Type-specimens.—Male holotype and nine male and one female para- types collected by R. M. Shelley, 9 August 1976, from viny undergrowth near Chiles Branch Creek, along S.C. Hwy. 48, 0.1 mi W junction S.C. Hwy. 63, edge of Sumter National Forest, 12 mi S Greenwood, Greenwood Co., S.C. (A1422). Male paratypes deposited in Florida State Collection of Arthropods, Zoologisch Museum Amsterdam, and personal collection of Richard L. Hoffman. Diagnosis.—Paranota red, with red stripe of varying width along an- terior edge of collum; sternal process between third pair of legs of males longer than width of adjacent coxae, bent anteriad distally, divided apically; gonopods with following characteristics: prefemoral process broadly curved ventrad, apically bifurcate, medial component longer than lateral; expansion of proximomedial edge of telopodite relatively long, irregularly notched; arc of telopodite relatively short and high; subapical lobes of telopodite moderate in size, distinct; solenomerite longer than lateral lobe; distal extremity of telopodite extending beyond outermost point in curvature of prefemoral process. Holotype —Length 46.0 mm; width of segment six 10.1 mm, of seg- ment ten 10.3 mm, of segment fifteen 9.6 mm; W/L ratio 22.4%; depth/ width ratio 58.3%. Color in life—Paranota red, metaterga black; broad red stripe along anterior edge of collum, connecting with paranotal spots; narrow red stripe along caudal edge of collum, faint red stripe along caudal edge of second metatergite. Epicranium, genae, and frons black, epicranial suture and clypeus lighter; antennae dark brown. Venter and legs generally light brown, sterna and basal podomeres of segments 6-15 darker. Structural details similar to those of C. catawba with following excep- tions: Width across genal apices slightly narrower, 4.6 mm; genae distinctly impressed. Paranota relatively flat. Peritremata more distinctly elevated, result- ing in blunt paranotal corners on segments 2-4. Ozopores conspicuous, opening dorsolaterad. Sternal process between third pair of legs (Fig. 13) bent anteriad dis- tally, longer than width of adjacent coxae; lobes between fourth pair of legs more prominent than in C. catawba; sternum of segment 6 broader VOLUME 90, NUMBER 2 313 9 10 Figs. 7-10. Telopodites of left male gonopods of Croatania spp. holotypes, lateral views. 7, C. catawba. 8, C. saluda. 9, C. simplex. 10, C. yemassee. 314 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON and more deeply recessed between seventh legs. Postgonopodal sterna less obviously bilobed, more platelike; subcoxal spines completely ab- sent. Coxal tubercles present on legs 18-25; prefemoral spines distinct on legs 7-30. Gonopods (Figs. 3, 8) not overlapping, extending forward just beyond anterior edge of aperture, inserting into depression of sternum between seventh pair of legs. Coxa smaller than in C. catawba. Prefemoral process not globose basally, broadly curved ventrad, bifurcate apically, medial component longer than lateral component, width at base of bifurcation subequal to that at base of process, tip extending beyond apex of telopodite. Expansion of proximomedial edge of telopodite longer and more irregularly notched than in C. catawba; telopodite more highly arched, distalmost point extending beyond outermost point in curvature of prefemoral pro- cess; subapical lobes reduced in size, more distinctly separated from solenomerite than in C. catawba, solenomerite longer than lateral lobe; ridge along inner face of telopodite low, indistinct, without projections; course of prostatic groove as in C. catawba. Male paratypes—The W/L ratios of the nine male paratypes range from 22.1%-24.6%, average 23.3%. All specimens display a red stripe of varying width along the anterior edge of the collum. Two individuals with narrow stripes also have narrow paranotal markings covering little more than the peritremata on the midbody segments. On the holotype and other paratypes, the red color covers the entire paranotum and extends inward along the caudal edge of the metaterga. On the gonopods, the degree of jaggedness of the proximomedial expansion of the telopodite varies, and no two are alike, not even on the same individual. The relative lengths of the two components of the prefemoral process also vary, al- though the medial one is always longer. Female paratype.—Length 40.0 mm, width 9.9 mm, W/L ratio 25.0%, depth/width ratio 70.0%. The specimen displays a broad red stripe along the anterior edge of the collum and. wide paranotal markings. The sternal process between the third pair of legs is as in C. catawba. On the cypho- pods, the membrane enlargement is smaller than that of C. catawba, and the receptacle, valves, and operculum are as in the type species. Variation Variation throughout the range of C. saluda is essentially like that discussed above for the paratypes. The length of the proximo- medial expansion varies and is shortest on a male from Edgefield Co., 11.3 mi NW Edgefield (A1418). The degree of separation of the soleno- merite and lateral lobe (as viewed medially ) is narrowest in specimens from Edgefield and McCormick counties, at the southern edge of the range. The length of the solenomerite relative to the lateral lobe varies, but the former is always longer. A few individuals have projections on the ridge carrying the prostatic groove, but these are generally absent. VOLUME 90, NUMBER 2 315 Ecology.—This species is abundant in August throughout its range and is usually encountered under thin layers of leaves near water sources. It was taken from predominantly pine forests and some that were pre- dominantly hardwood. The type specimens were collected from an area covered with honeysuckle (Lonicera sp.) and low-growing vines, a habi- tat that is rarely investigated by diplopod collectors. Distribution —Croatania saluda is known from the central and south- central Piedmont of South Carolina, from Newberry Co. to the Savannah River (Fig. 17). Specimens have been examined as follows: South Carolina: Newberry Co., 11.7 mi NE Newberry, along S.C. Hwy. 81 at Enoree R., 2¢, 42, 5 August 1976, R. M. Shelley (A1419); 9.3 mi NE Newberry, Mollys Rock Picnic Area, Sumter National Forest, 4¢, 32, 5 August 1976, R. M. Shelley (A1424); and 9.4 mi NW Newberry, along S.C. Hwy. 32 at Indian Cr., 116, 42, 5 August 1976, R. M. Shelley (A1420). Greenwood Co., 4.2 mi E Ninety Six, near Wilson Cr., 146, 21 May 1960, L. Hubricht (RLH); 12 mi S Greenwood, viny undergrowth area along Chiles Branch Cr., along S.C. Hwy. 48, 0.1 mi W ject. S.C. Hwy. 63, edge of Sumter National Forest, 106, 2, 9 August 1976, R. M. Shelley (A1422) TYPE-LOCALITY; and 12 S Greenwood, unnumbered rd. at Cuffytown Cr., edge of Sumter National Forest, 26, 3°, 9 August 1976, R. M. Shelley (A1423). Edgefield Co., 11.3 mi NW Edgefield, along S.C. Hwy. 283 at Tur- key Cr. near McCormick Co. line, 86 62, 8 August 1976, R. M. Shelley (A1418); and 11.4 mi W Edgefield along S.C. Hwy. 68, 0.2 mi N jet. S.C. Hwy. 52, 6, 42, 8 August 1976, R. M. Shelley (A1421). McCormick Co., 4 mi E McCormick, along U.S. Hwy. 378 at Hard Labor Cr., 6, 2, 8 August 1976, R. M. Shelley (A1425). Remarks.—The gonopod of C. saluda is superficially similar to that of C. catawba but differs in several important features, the most obvious being the prefemoral process, which is apically bifurcate in the former and simple in the latter. This structure is also more broadly curved in C. saluda and much thicker (subglobose) basally in C. catawba. On the telopodite, the expansion of the proximomedial edge is longer and more irregularly notched in C. saluda and subequal in length to the prefemoral process on some individuals. In the type species the expansion is always shorter than the length of the prefemoral process and usually consists of two main groups of notches, one at either end. The arc formed by the curvature of the telopodite is longer and broader in C. catawba, although this difference is not readily apparent in the holotypes. The subapical lobes are much larger in C. catawba, but the solenomerite is more dis- tinctly separated from the lateral lobe in C. saluda. In the latter species the solenomerite is longer than the lateral lobe, whereas in C. catawba they are either subequal, or the solenomerite is shorter. The two species can also be distinguished by non-sexual characters, and 316 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON the yellow and red paranotal colors afford a ready basis for field identifica- tions. The paranotal corners of segments 2-4 are more blunt in C. saluda due to thicker peritremata on these segments. Ventrally, the sternal pro- cesses on the fourth segments are similar in both species, but the post- gonopodal sterna are flatter and more platelike in C. saluda. The de- pression in the sternum between the seventh legs is deeper and broader in C. saluda. Croatania simplex, new species Figs. 4-5, 9, 14 Type-specimens.—Male holotype (A1417) and fourteen male and seven female paratypes collected by R. M. Shelley, 4 August 1976, from Chester State Park, Chester Co., S.C. Male and female paratypes deposited in Florida State Collection of Arthropods, Zoologisch Museum Amsterdam, and personal collection of Richard L. Hoffman. Diagnosis.—Paranota yellow, with or without yellow stripe along an- terior edge of collum; sternal process between third pair of legs of males shorter than width of adjacent coxae, straight, divided at midlength; gonopods with following characteristics: prefemoral process short, ter- minating well below level of tip of telopodite, usually of subequal thickness and without apical bifurcation; expansion of proximomedial edge of telopo- dite relatively shallowly notched; subapical lobes of telopodite greatly re- duced, indistinct; solenomerite elongate and broadly curved toward center of arc of telopodite, acuminate, much longer than lateral lobe; distal ex- tremity of telopodite extending slightly beyond outermost point in curva- ture of prefemoral process. Holotype.—Length 43.7 mm; width of segment six 10.6 mm, of segment ten 10.7 mm, of segment fifteen 10.2 mm; W/L ratio 24.5%; depth/width ratio 57.0%. Color in life——Paranota bright lemon yellow, metaterga black; broad yellow stripe along anterior edge of collum, connecting with paranotal spots. Epicranium and interantennal region black, genae and clypeus lighter; antennae brown. Venter and legs essentially uniform in color, lighter than paranota. Structural details similar to those of C. catawba with following excep- tions: Width across genal apices slightly narrower, 5.0 mm; genae distinctly impressed. Paranota relatively flat. Peritremata slightly more distinct on segments 2-4, resulting in less rounded paranotal corners. Sternal process between third pair of legs (Fig. 14) divided at mid- length, leaning anteriad but straight, not bent distally, shorter than width VOLUME 90, NUMBER 2 317 Figs. 11-15. Median sternal processes of fourth segments (between third pairs of legs) of Croatania spp., caudal views. 11-12, C. catawha; 11, Holotype; 12, Female paratype. 13, C. saluda, holotype. 14, C. simplex, holotype. 15, C. yemassee, holotype. of adjacent coxae; sternum more deeply and broadly recessed between seventh legs; postgonopodal sterna less obviously bilobed, more plate- like, subcoxal spines absent except for small pair on segment 8. Coxal tubercles present on legs 13-27; prefemoral spines distinct on legs 5-30. Gonopods (Figs. 4, 9) overlapping at distal curvature, extending beyond anterior edge of aperture and inserting into indentation of sternum be- 318 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON tween seventh legs. Prefemoral process greatly reduced, not globose basally, broadly curved ventrad, terminating well below level of tip of telopodite. Expansion of proximomedial edge of telopodite with shallow notches; distal curvature of telopodite slightly higher than in C. catawba, distal- most point extending slightly beyond outermost point in curvature of pre- femoral process; subapical lobes greatly reduced, inconspicuous, medial lobe almost nonexistent, lateral lobe slightly larger; solenomerite elongate and broadly curved, acuminate, extending inward beyond tip of pre- femoral process, much longer than lateral lobe; ridge along inner face of telopodite indistinct, without projections, course of prostatic groove as in C. catawba. Male paratypes——The W/L ratios of the fourteen male paratypes range from 21.5%-24.5%, average 22.8%. Three individuals lack the stripe along the anterior edge of the collum. Two have an orange tint to the paranotal markings which extend inward on one suggesting a stripe along the caudal edge of the metaterga. On the gonopods, the prefemoral processes vary in several respects. Some are broadly curved, whereas others are relatively flat. The structure is apically bifurcate on two specimens, with medial component longer and more prominent than the lateral one (Fig. 5). It is subglobose basally on others. The subapical lobes, especially the medial one, are more pronounced on most paratypes than on the holotype, and the solenomerite is slightly bisinuate on some specimens. Female paratypes—The seven female paratypes have a higher W/L ratio than the males, the average being 24.6%. Two individuals lack the yellow stripe along the anterior edge of the collum. The sternal process between the third pair of legs is reduced from the condition in C. catawba. On the cyphopods, the membrane enlargement is smaller than that of C. catawba, and the receptacle, valves, and operculum are as in the type species. Ecology.—This species is extremely abundant at the type-locality. Most individuals were found under thin leaf cover in predominantly hardwood areas on slopes near creeks or along the edges of paths, and were taken in association with Sigmoria latior intergrades (see Shelley 1976). Distribution.—Known only from the type-locality. Remarks.—Croatania simplex is distinct from its congeners in the greatly reduced subapical lobes of the telopodite, elongate solenomerite, and short prefemoral process. The sternal process of the fourth segment is also unique, being shorter than the margins of the adjacent coxae and divided for a greater proportion of its length, beginning at midpoint. The species resembles C. catawhba in the yellow paranotal color and the ten- dency of the prefemoral process to be subglobose basally. It is similar to C. saluda in the arc of the telopodite, the solenomerite’s being longer VOLUME 90, NUMBER 2 319 than the lateral lobe, and the tendency of the prefemoral process to be bifurcate. Croatania yemassee, new species Bigs, 64 10 1s Type-specimens.—Male holotype collected by L. Hubricht, 20 September 1959, from low oak woods, 8 mi S Hardeeville, Jasper Co., S.C. (RLH). Male and female paratypes collected by L. Hubricht, 5 July 1959, from a small island 7 mi E Frogmore, Beaufort Co., S.C. (RLH). Diagnosis.—Sternal process between third pair of legs of males straight, subequal to or slightly longer than width of adjacent coxae, divided apically; gonopods with following characteristics: prefemoral process bisinuately curved, bent sharply ventrad near acuminate tip; expansion of proximo- medial edge of telopodite reduced, with only small denticulations; arc of telopodite short and high; subapical lobes reduced, indistinct; distal ex- tremity of telopodite extending beyond outermost point in curvature of prefemoral process. Holotype.—Length 40.9 mm; width of segment six 10.5 mm, of segment ten 10.7 mm, of segment fifteen 10.4 mm; W/L ratio 26.2%; depth/width ratio 46.7%. Color in life unknown, evidence of stripe along anterior edge of collum in preserved specimen. Structural details similar to those of C. catawba with following excep- tions: Width across genal apices slightly narrower, 4.7 mm. Antennae longer, extending slightly beyond caudal edge of third metatergite. Genae with medial impression. Frontal setae not detected and presumed absent. Peritremata more distinctly elevated, resulting in blunt paranotal cor- ners on segments 2-4. Ozopores more conspicuous, opening dorsolaterad. Sternal processes between third pair of legs (Fig. 15) straight, not bent anteriad distally, divided apically, slightly longer than width of adjacent coxae; lobes between fourth pair of legs more prominent than in C. catawba; sternum between seventh legs deeply recessed. Postgonopodal sterna less obviously bilobed, more platelike; subcoxal spines subtending only posterior legs on segment 8, absent thereafter. Coxal tubercles present on legs 8-27; prefemoral spines distinct on legs 5-30. Gonopods (Figs. 6, 10) not overlapping, extending forward just beyond an- terior edge of aperture. Prefemoral process upright, bisinuately curved, not globose basally, bent ventrad apically and narrowing abruptly to acuminate tip, tip directed toward midpoint of telopodite. Expansion of proximo- medial edge of telopodite reduced, with only small denticulations; telopo- dite highly arched and bent abruptly dorsad, distal extremity extending beyond outermost point in curvature of prefemoral process; subapical 320 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 16. Left cyphopod of female paratype of C. catawba, ventral view, in situ. me, membrane enlargement; r, receptacle; v, valves. lobes reduced, inconspicuous, lateral lobe subquadrate, serrate; soleno- merite blunt, at approximately same level as distal portion of prefemoral process; ridge along inner face of telopodite not distinctly raised, with basal projection; course of prostatic groove as in C. catawba. Male paratype.—The male paratype is badly fragmented and cannot be measured but agrees essentially with the holotype in all structural details. On the gonopods, the prefemoral process is more pointed, and the ex- pansion of the proximomedial edge of the telopodite is more distinctly notched. Female paratype.—Length 44.6 mm, greatest width 11.4 mm, W/L ratio 25.6%, depth/width ratio 66.7%. Color in life unknown, but there is evi- dence of a stripe along the anterior edge of the collum. Frontal setae are apparently absent, and the sternal process between the third pair of legs is as in C. catawba. On the cyphopods, the membrane enlargement is re- duced over the condition in C. catawba, and the receptacle, valves, and operculum are as in the type-species. Distribution.—Coastal islands and outer Coastal Plain of southern South Carolina (Fig. 17). Specimens have been examined as follows: South Carolina: Jasper Co., low oak woods, 8 mi S Hardeeville, ¢, 20 VOLUME 90, NUMBER 2 321 September 1959, L. Hubricht (RLH) TYPE-LOCALITY. Beaufort Co., small island 7 mi E Frogmore, 3, 2, 5 July 1959, L. Hubricht (RLH). Remarks.—Croatania yemassee is distinct from its congeners in the ap- parent loss of frontal setae, the bisinuately curved prefemoral process, and reduction in the expansion of the proximomedial edge of the telopodite. The depression in the sternum between the seventh legs is also much deeper than in the other species. Other than the diagnostic characters of the genus, it has little in common with C. catawba but is similar to C. saluda in the height and breadth of the arc of the telopodite, the relatively flattened postgonopodal sterna, and the blunt paranota of segments 2-4. The reduc- tion in the subapical lobes of the telopodite, a feature shared with C. sim- plex, is an example of convergence and not taxonomically significant. Ecology A striking ecological characteristic of Croatania is the abundance of its species in July and August, the hottest, driest, months of the year. The millipeds seem to thrive in such conditions, when many other species are scarce (see Shelley, 1976, for a comment on the effects of climate on Sigmoria latior). Other diplopods that would be expected in the Piedmont, such as species of Cambala, Polyzonium, Pseudopolydesmus, Ptyoiulus, and Scytonotus, were notably absent in August 1976, presumably because of the harsh conditions. Croatania catawba and C. saluda were frequently found in association with Pachydesmus crassicutis incursus, which is com- mon in the same habitats (Shelley and Filka 1977). In addition to these xystodesmids, Sigmoria latior intergrades, S. l. hoffmani Shelley, and two undescribed forms currently under study, the only diplopods that were even moderately abundant in the Carolina Piedmont in August 1976 were Narceus americanus (Beauvois) and Abacion magnum (Loomis). Croatania is also unusual in the variety of habitats it occupies. In the Kings Mountain region it occurs in climax deciduous forests with thick leat layers, traditional xystodesmid habitat. Outside this region, however, it in- habits intermediate successional stages of mixed pine-hardwood and was found in some localities dominated by pine. At Mollys Rock Picnic Area, Newberry Co., South Carolina, for example, several individuals of C. saluda were discovered under pine bark mulch on a walkway through a pine forest. At the type-locality of C. saluda, individuals were found in a pine area covered with honeysuckle and low-growing vines, a habitat that would usually be bypassed by milliped collectors. These two localities were some- what atypical, and in most places Croatania occurred under thin leat layers relatively close to water sources. Sites with moderate slopes lead- ing to rivers or creeks usually produced both Croatania and Pachydesmus at the bases of the slopes. As with P. c. incursus (Shelley and Filka ey) bo bo PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 1977), specimens of Croatania from the Kings Mountain region are notably larger and more robust than those from areas farther south with a greater proportion of pine, probably a reflection of more favorable environmental conditions in this region. Distribution The distribution of Croatania (Fig. 17) is of interest in the absence of obvious geographical or ecological interpretations. Discounting C. yemas- see, which is isolated on the southeastern coast of South Carolina, the ranges of the other three species in the state cannot be correlated with any factor; they all occur in the same general area of the central Piedmont and in essentially the same habitats. The Savannah River is perhaps the southern boundary for C. saluda (to be determined by future field work in Georgia), but there is no ready geographical explanation for its apparent northern limit. It was collected as far north as the Enoree River, but its range is bissected by the much larger Saluda River. Likewise, C. catawba was collected as far south as the Tyger River, but its range is bissected by the much larger Broad River. Current data indicate that the ranges of these two species abut near the Enoree and Tyger Rivers and the Union-New- berry Co., S.C., line. Further collecting may show that they overlap. Since the ranges of both species span several relatively large rivers, it seems that stream capture and resultant changes in drainage boundaries may have been important factors in determining their present distributions. Com- ment on C. simplex, known from only one locality, is deferred until its complete range can be more precisely defined. The distribution of C. catawba in North Carolina can perhaps be ex- plained geographically. According to Stuckey (1965), the Kings Mountain range of Cleveland and Gaston counties continues northward through a series of hills and ridges to Anderson’s Mountain in Catawba Co., just north of Lincoln Co. Perhaps the distribution of C. catawba in North Carolina is associated with this physiographic feature, and if so, it should occur in Catawba Co. and the northern part of Gaston Co. Efforts to find the milliped in these areas have so far been unsuccessful but remain an objective of future studies. Relationships Generic—The presence of large sternal processes on the fourth seg- ments, bilobed postgonopodal sterna, and similarities in the distal portions of the telopodites, indicate a close relationship between Croatania and Cleptoria. The former is a more advanced, specialized genus as evidenced by the more highly developed prefemoral process, ornate proximomedial expansion of the telopodite, and subapical lobes. In Cleptoria the pre- VOLUME 90, NUMBER 2 323 rf f Po, *s 0°% . ° A .* ~ root. : t : 4G 52% “e \ ° e v* bone. te cce coos e ooo coe z A 3 *.f2* ; ‘ \\ *s is =) 2 2 ROG O0CG od s ae Fi oa Sor iee owe ace : : : s : 2 Aye © wa%e e “ee ra oe ° ‘S ° er | 3 Z : % 2 i a z “. e Ns °e, : ° 5 ; voce os ooctee S ie g “ 4 . 8 Re ae Pate feast £ oh, ey ope a . °° Scherer ere ness mran rg : : : Zacks ate i ea te ede GS : “eo % OCS RO mane ys 2 we 8 r ese cee”, say FZ “. its 2 O- r 5 CROOK re ry ©. Ac cceee Sacboves er og “75 Y H °. i 4 “3, By one? Se) : Peccece y 3. ° e es “e, ~ S ys Peer es yt ae) We 2 pe b % 5 . - Wy (ome gente see a Pa 3 Ds Sy : OF O t 7 *e e *. 3 is ‘ os, * “ a ie YH 0, ¢ tR0000CS Py eee” ‘ee oeo o%, ° is a ot =. 5 $ © > 0 Sag ere Z Sey is >: @4@6.: a *. S $ “oe 2 an) 2 3 ey occeee o% 5000" 8% Seyi ys : x . oneces Le e ie Ont PYG. of rs 2 “e é r} : m7 BH ° ene af ° . ote > Ae et ee e coe Se® som rs ° ° “o 2) yy Ie - ry oo more yy os Sh . & 5 Pene* %., *og 2 ? e Woeters ate trae a 4 C. catawba e@ sth a COS ao =— — PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 330 ‘OFT UL poydeisojoyg “9/61 “O1S9N VOLUME 90, NUMBER 2 331 with slightly convex and nearly horizontal ventral profile of caudal peduncle. Distance between snout tip and pectoral-fin origin 27.3 (27.3- 29.7, x = 28.5) [29.1-30.4, x = 29.8]. Distance between snout tip and pelvic-fin origin 43.7 (43.7-46.3, x = 44.9) [46.8-50.1, x = 48.4]. Dis- tance between snout tip and anal-fin origin 57.3 (56.5-59.5, « = 57.7) [56.9- bree —o0 | Gaudal) peduncle depth 10:8 (103-112, x = 10:77) [10 1— 10.7, x = 10.3]. Caudal peduncle length 10.8 (10.4-11.8, = 11.3) [9.8-11.8, os ieSe Head relatively short and deep. Head length 27.0 (26.1-27.7, « = 26.9) [27.0-29.6, x = 28.6]. Snout blunt and lower jaw protruding beyond upper jaw. Mouth gape angled dorsally. Posterior ventral border of maxillary reaching or nearly reaching a point vertically below anterior border of iris of eye. Horizontal eye diameter 9.8 (9.8-12.1, x = 10.9) [9.6-13.1, « = REG FRoWOueenetin 7.7 (7.0-6.5, % =-7.8)) [7.8-8.5, x = 8.2]. east bony interorbital width 8.5 (8.3-8.7, « = 8.5) [8.3-8.8, x = 8.7]. Dorsal-fin rays ii,9 in all specimens (last ray not split to its base). Dorsal-fin length (= length of longest ray) 33.9 (30.4-34.4, * = 32.3) [29.1-30.8, « = 30.2]. Distal margin of dorsal fin pointed in male, rounded in female. Adipose fin present. Anal-fin rays iv,28 (iv,28 in three, iv,29 in one, and iv,30 in two males) [iv,27 in one, iv,28 in three females]. Last anal-fin ray split to its base. Anal-fin length (= length of longest ray, secondmorancheal ray) 22.) (181-209) x = 19:9) [18:7—21.5, « = 20-5). Margin of anal fin slightly convex to slightly concave. Pectoral-fin rays i,12 (i,12 in six and i,13 in one male) [i,12 in three and i,13 in one female]. Posterior tip of longest pectoral-fin ray reaching beyond pelvic-fin origin. Rectoralnnmplemeth 24:4 (22.9-945, x = 23.8 n = 5) [21.6-23.6, x = 22.6]. Pelvic fin usually reaching posterior to anal-fin origin in mature males and not quite to that origin in females. Pelvic-fin rays i,7 in all specimens. Pelvic-fin length 20.8 (18.6-20.8, x = 19.7) [14.5-19.0, x = 17.6]. Caudal fin 10/9, its dorsal and ventral lobes equal in length. Small bony hooks present on distal one-half to one-third of fourth unbranched anal-tin ray, one hook per segment except absent on distal 2 to 4 segments. Similar hooks on branched portions of first (anterior) to eighth or tenth branched anal-fin rays. Hooks most common on posterior branch of each fin ray where they occur on each ray segment except for distal 2 to 4 or 5 seg- ments. Anterior branch of each fin ray with an occasional hook. Dorsal fin with second unbranched ray with distally located hooks distributed similar to those of fourth unbranched anal-fin ray. First through third branched rays of dorsal fin bearing small hooks with a distribution similar to those on branched anal-fin rays. Scales cycloid. Lateral line incomplete; perforated lateral-line scales 10 (7 in one, 8 in one, 9 in two, 10 in five, and 13 in one specimen). Scales ‘OFT UL poydeisojoyg ‘9/6[ ‘O1SeN ONY ‘sojooleg “seuozeuly JO 97e]S * ‘od Aje.ied “TSG wut Gg s[eulof ‘ZTO9OTS INNSA ‘Setoads Mou “2foj090s woohugossaydAyT *¢ “BI Z S Ho 0 ‘a es WN q 5 = © va - al Oo } WN 4 << v oe o a S pO ou ro fH cy ° © 4 A ea eal © } ac = VOLUME 90, NUMBER 2 333 in a lateral series 33 (31 in one, 32 in three, 33 in five, and 34 in one specimen). Scale rows between lateral line and dorsal-fin origin 7 in all specimens. Scale rows between lateral line and anal-fin origin 5 in all specimens. Scale rows around caudal peduncle 14 in all specimens. Pre- dorsal scales 10 in all specimens. Teeth tricuspid. Premaxillary with 2 (2 in nine and 3 in one) teeth in outer row and 6 (5 in one, 6 in six, and 7 in three specimens) in inner row. Maxillary with 10 (3 in one, 4 in four, 5 in three, 6 in one, and 10 in one specimen). Dentary with 5 large teeth followed by 6 small teeth (4-4 in one, 4—5 in two, 5-5 in one, 5-6 in three, 5-7 in one, and 6-10 in one specimen ). Total vertebrae including Weberian apparatus and terminal half centrum 32 (32 in nine and 33 in one). Gill rakers 9/20 in one specimen, 9/17 in another. Color in alcohol.—(Description from holotype after about 2 weeks in for- malin and 6 months in alcohol, Fig. 1). Entire body dusky brown with a dark vertically elongate “shoulder” spot immediately posterior to operculum ex- tending from near pectoral-fin origin dorsally to about 2 horizontal scale rows above lateral line. Body area posterior to this nearly vertical bar paler than remainder of body, including belly (silvery pigment destroyed by initial preservation in formalin). Posterior to pale area, body again dark but comparatively pale posterior to body cavity. A broad, rather poorly de- fined, dark brown horizontal stripe extends from dorsal region of vertical dark area described above onto caudal peduncle to caudal-fin root. Back above about third horizontal scale row darker than general body color. Top of back anterior to dorsal fin and top of head very dark, similar to a diffuse dark narrow stripe on back along back posterior to dorsal-fin base and extending to dorsal portion of caudal-fin root. Head about same dusky color as back. Opercular and orbital region ventral to eye paler than remainder of head. Scales along base of anal fin very dark, produc- ing a nearly black dusky base to anal fin. Immediately distal to this area base of anal fin nearly white in recently preserved specimens. In speci- mens preserved for over 1 year white may be absent, leaving this area of fin dusky hyaline. Anterior portion of white of anal fin not extending dis- tally more than half of length of fin rays and usually extending distally less than about one-third of fin-ray length. Distal to white stripe, anal fin hyaline except for a heavy “dusting” of dark chromatophores which is especially dark on about 8 anteriormost rays. Pectoral, pelvic, and caudal fins hyaline except for a fairly dusky appearance produced by scattered dark chromatophores. Dorsal fin with same color except for a large black spot covering nearly two-thirds to four-fifths of distal distance of third to about sixth fin rays and their connecting membranes. Distal halt 334 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON of second dorsal-fin ray white as are distal tips of third, fourth, and fifth fin rays. Adipose fin dusky. Color in life—Brown to black pigment about same as described above under color in alcohol, but much silvery pigment in area of abdominal cavity, on head below eye, and on lower jaw. Both sexes with a “warm” but relatively pale chocolate brown color with an orange-red aspect modi- fied by pale blue and green producing a pale violet appearance. Colors other than brown rather highly reflectant and produced by densely scat- tered small blue, blue-green, red, orange, and gold chromatophores over entire body. Gold, blue, and green chromatophores especially common on ventral half of body. Back considerably darker than remainder of body. A pink to deep-red spot (sometimes fully as brilliant as red in eye) just dorsal to fifth to seventh lateral-line scales. This spot may cover area of 2 or 3 scales. A similar red color on posterior border of caudal peduncle but this mostly obscured by brown to black chromatophores. Just anterior to eye and posterior to dorsal one-third of premaxillary is a vertically elongate metallic copper colored bar. Top of head dark brown. Opercle pale bluish to greenish silvery color with scattered brown to black chromatophores. Sometimes scattered pale red chromatophores occur over ventral opercular area. Lower jaw silvery white. Tip of lower jaw and dorsal third of premaxillary dusky brown. Just posterior to dorsoposterior portion of eye occurs a reflectant gold spot (often partly ob- scured by brown chromatophores) about size of pupil of eye. Fins dusky hyaline. Caudal fin and pectoral fin without other pig- ment. White of anal fin (distributed as described above under color in alcohol) with an intense milky appearance. In females milky-white may disappear posteriorly by about tenth to twelfth fin rays whereas in males it may be present posteriorly as far as posterior terminal fin ray. Scat- tered golden-orange chromatophores occur over remaining parts of anal fin. Distal tips of anal-fin rays with membrane between them nearly black. Pelvic fin with red and orange chromatophores along its basal one-third to one half; distally hyaline. Adipose fin colorless except for scattered dark brown or black chromatophores. Dark spot of dorsal fin distributed as described under color in alcohol. Spot black in life, sur- rounded basally and anterodistally by white pigment in males which may also contain some orange pigment. In adult females distal “white” area of dorsal fin often a deep intense milky orange color. Some of this orange pigment also occurs in basal “white” areas of dorsal fin in females. Dorsal half of eye (exclusive of pupil) deep red except for black area immediately dorsal to pupil. Ventral half of eye dusky slivery often with a darker dusky area immediately ventral to pupil. Eye often with appearance of a vertical bar through its center. VOLUME 90, NUMBER 2 335 Hyphessobrycon erythrostigma (Fowler ) Figs. 4-8 Hemigrammus erythrostigma Fowler, 1943:33, original description, figure, “Brazil,” holotype only, aquarium specimen. Hyphessobrycon rubrostigma Hoedeman, 1956:312, original description, fig- ure, “Colombia, aquarium specimens.—Gery, 1965:22, Igarapé Préto, Brazil, 60 km downstream from the border zone between Peru, Colombia and Brazil. Hyphessobrycon erythrostigma Sterba, 1970:98, aquarium description and color photograph of adult pair, use of nomenclatural combination credited to Gery in litt—Geéry, 1972:7, placed H. rubrostigma in synonymy of H. erythrostigma. Material examined.—Holotype, ANSP 70208, 46.4 mm SL, female bearing eggs, aquarium specimen said to be imported from Brazil—USNM 216613, 8 (43.2-60.6 mm SL), imported aquarium specimens without certain lo- cality but presumably exported from Leticia, Colombia, imported into United States during 1971-1976—USNM 174944, 6 (29.0-40.8 mm SL) (an additional 7 juvenile specimens not examined), aquarium specimens without certain locality, said to be from Peru, imported 1958. The above material is not satisfactory in terms of locality information but apparently represents the best available. It does have the merit of exhibiting growth changes in morphological characters, especially those associated with the attainment of maturity in the males and therefore provides a guide for future examination of population samples with locality data. Diagnosis.—This species may be distinguished from all members of the genera Hyphessobrycon, Hemigrammus or related genera by the same char- acters as listed for H. socolofi above except that it differs from that species by the following characters. Both sexes with white pigment along base of anal fin extending 75-90% of distal distance onto its anterior lobe (extending no further than about 50% out onto anterior anal-fin lobe in H. socolofi). compare Figs. 2 and 3 of H. socolofi with Figs. 5 and 6 of H. erythrostigma. Vertebrae 33 in all specimens examined (in H. socolofi vertebrae 32, one specimen with 33). Mature adults about 40-60 mm SL (in H. socolofi mature adults reach about 30 to nearly 40 mm SL). Length of adults should be examined carefully in wild population samples since it is pos- sible in aquaria to obtain “stunted” adults of H. erythrostigma at 30-40 mm SL. Males nearly without small bony hooks in dorsal and anal fins (very old specimens with small, rounded bony processes on theses fin rays) (numerous small bony hooks present in dorsal and anal fins of H. socolofi). Mature males with anterior second, third, and fourth dorsal-tin rays about 38-50% of SL (about 30-35% in H. socolofi). Mature males with fourth and fifth anal-fin rays about 27-35% of standard length (about 336 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 18-22% in H. socolofi). Both sexes with snout 6.9-7.9% of standard length (7.8-8.5% in H. socolofi). Description—Body deep, sides compressed, greatest depth at anterior base (origin) of dorsal fin, 51.1 (36.1-48.2, x = 43.2) [36.2-53.5, x = 45.6]. Predorsal body profile gently convex but concave at nape and again con- vex at snout. Body profile along base of dorsal fin straight to slightly convex, nearly straight to very slightly convex between dorsal fin and adipose fin, concove in region of caudal peduncle. Dorsal fin nearer to snout tip than to caudal-fin base. Distance between snout tip and anterior dorsal-fin origin 53.5 (44.7-49.1, x = 47.0) [45.9-53.5, « = 48.8]; distance between anterior dorsal-fin origin and caudal-fin base 61.2 (56.0-64.7, x = 61.0) [54.5-62.5, x = 59.2]. Distance between posterior border of eye and dorsal-fin origin as a percent of distance between dorsal-fin origin and caudal-fin base 62.3 (52.9-58.8, « = 54.8) [53.2-62.3, x = 58.0]. Ventral body profile moderately convex in both sexes between symphysis of lower jaw and anal-fin origin. Body profile strongly convex at origin of anal fin, then nearly straight to gently convex to basal termination of anal fin. Ventral profile of caudal peduncle slightly concave. Distance between snout tip and pectoral-fin origin ? (26.7-30.2, * = 28.5) [27.7-30.7, x = 29.8, n = 6]. Distance between snout tip and pelvic-fin origin ? (40.9-44.5, « = 492.6) [44.3-48.2, x = 45.6, n = 6]. Distance between snout tip and anal-fin origin P (52.0-59.8, « = 56.1) [56.6-64.7, x = 59.9, n = 6]. Caudle peduncle depth 9.9 (9.2-11.1, « = 9.9) [9.6-11.1, = 10.3]. Caudal peduncle length IULicy (UES SO) se = 0) [Sica ce == ULI] Head relatively deep and short. Head length 27.8 (26.8-28.1, x = 27.8) [26.2-29.9, x = 28.1]. Snout blunt and lower jaw protruding beyond upper jaw. Mouth gape angled dorsally. Posterior ventral border of maxillary reaching to or somewhat beyond a vertical line drawn ventrally from anterior border of eye. Horizontal eye diameter 11.6 (9.2-14.0, x = 11.7) [9.8-13.0, x = 11.7]. Snout 7.3 (7.0816, x = 7.6) (G9 Seo areal Least bony interorbital width 8.8 (7.7-9.0, = 8.3) [7.8-8.8, x = 8.5]. Dorsal-fin rays ii,9 in all specimens (last ray not split to its base). Dorsal- fin length (= length of Jongest ray) ? (25.4505, x = 37a eo se0) * = 31.5, n = 6]. Distal margin of dorsal fin pointed in male, rounded in female. Adipose fin present. Anal-fin rays iv,29 (iv,27 in two, iv,28 in three, iv,29 in two males) [iv,28 in two and iv,29 in five females]. Last anal-fin ray split to its base. Anal-fin length (= length of longest ray, which is either last unbranched or first branched ray) ? (21.9-35.7, x = 29.1) [19.7- 21.0, x = 20.2, n = 6]. Posterior margin of anal fin (beyond anterior lobe) slightly convex but strongly concave anteriorly where it approaches and is continuous with anterior lobe. Pectoral-fin rays i,12 (i,12 in one i,13 in six males) [i,12 in six, i13 in one female]. Posterior tip of longest pectoral- fin ray reaching beyond pelvic-fin origin. Pectoral-fin length ? (22.0-25.9, c ‘OL6I LIMO) DUdYsoLyjhisa uUuorhsgossaydhy, “y BA 337 PIQUIO[OT) “BIOQaT JO SoM 90uRy ; ‘CT99TS INNSN “( A “SIp owos, “SIP oulOs S = © Z, — en WN ae = ty ° 2 = 3S 6 N Ss = = @ fe) H 2 —Q fx] ze icon! Ly O WN O g A ea) ica Ss 6 rg a VOLUME 90, NUMBER 2 339 x = 23.7) [20.7-24.3, x = 22.5, n = 6]. Distal pelvic-fin tip usually reach- ing posterior to anterior anal-fin origin in males and not quite to or just to that origin in juveniles and adult females. Pelvic-fin rays i,7 in all spcemenss Eelvic-tin length ? (15.2-28.5, x = 22.3) 16.3-20.0, x = 18.24, n = 6]. Caudal fin 10/9, its dorsal and ventral lobes equal in length. Bony hooks nearly absent on dorsal and anal fins. Adult males may oc- casionally have one or two small hooks per fin ray on dorsal and anal fins. When present these hooks distributed among same numbered rays as in males of H. socolofi. Two senile males 56.7 and 60.6 mm SL with slight, rounded bony protuberances on fourth unbranched ray through to about eighth branched anal-fin rays and on second unbranched through second branched dorsal-fin rays. Scales cycloid. Lateral-line incomplete; perforated lateral-line scales ? (9 in two, 11 in three, and 14 in one specimen) [9 in one, 10 in three, and 11 in two specimens]. Scales in a lateral series 34 (33 in one, 34 in twelve, and 35 in one specimen). Scale rows between lateral line and dorsal-fin origin 7 in all specimens. Scale rows between lateral line and anal-fin origin 5 in all specimens. Scale rows around caudal peduncle 14 in all specimens. Predorsal scales 10 in all specimens. In scale counts above, those for holotype were taken from scale pockets and therefore in this specimen perforated lateral-line scales could not be counted and scale rows above and below lateral line not absolutely certain. Teeth tricuspid. Premaxillary with 3 (1 in one, 2 in five, and 3 in seven specimens) teeth in outer row and 6 (5 in five, 6 in eight, and 7 in one specimen) teeth in inner row. Maxillary teeth variable, 7 (1 in two, 2 in two, 3 in two, 4 in four, 5 in one, 6 in one, and 7 in one specimen). Dentary with 5 (4 in five and 5 in seven specimens) large teeth followed by 7 (5 in one, 6 in four, 7 in two, 8 in five, and 10 in one specimen) small teeth along sides of jaw. Total vertebrae including Weberian apparatus and terminal half cen- trum 33 in all specimens. Gill rakers 8/16 in one specimen. Color in alcohol.—Description from a male 43.2 mm SL (unless other- wise noted) initially preserved in 10% formalin and in alcohol for about six months, Figs. 4 and 5. Entire body dusky brown with dark “shoulder” spot beginning immediately posterior to operculum at a point just dorsal to a level equal in horizontal position to that of ventral border of eye and extending dorsally to about two horizontal scale rows above lateral line. In life, chromatophores of shoulder spot may extend ventrally below level of eye reaching near origin of pectoral fin (see Fig. 5). Area posterior to shoulder spot, paler than remainder of body including belly. Pale region equals area of red spot in life. Posterior to pale area body again dark, sometimes as dark as shoulder spot. This dark area fades posterior to region of body cavity. A broad, rather diffuse, dark brown horizontal stripe ex- ‘OFT UL poydeisojoyg “9/6 ‘eIquIojOD ‘eIoljeT JO JsaM _90uUe} -sIp oulos,, Ajjuotedde ynq UMOUYUN APIO] “TS wu g-Eg s[vulsF “ST9OTS INNSN “(49[M0y ) Dwsysolyzhva uoohsgossaydhy °9 ‘B14 Zz © x 1S) J ao N < S (xy e) > fH a O © nN — < Lv © e) | © faa) ica eer EH fy 'e) N O ‘a OQ fx] eal CO © jaa fall VOLUME 90, NUMBER 2 unknown. ality ~ ~ = — a — ~ — ow ~ =~ ~ — =~ ~ = — a 7 — — = = = —_— “5 — — e ~ — = =) = vo — ~~ — oy | =~ — A — a oma (Fowler), lp Gi, Hyphessobrycon erythrost SS \ QQ’ ‘ . we s 342 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON tends from dorsal region of pale area onto caudal peduncle to caudal-fin root. This poorly defined and often absent (especially in females) stripe occurs just at and dorsal to junction of hypaxial and epaxial myomeres. Back above about third horizontal scale row slightly darker than general body color. Top of back anterior to dorsal-fin and top of head very dark, similar to a very dark diffuse stripe on back along and posterior to dorsal- fin base and extending to dorsal portion of caudal-fin root. Dorsal half of head about same dusky color as back ventral to dark diffuse median stripe. Opercular and orbital region ventral to eye paler than remainder of head. Scales along base of anal fin dark dusky brown with pigment diffusely scat- tered. Immediately distal to this area base of anal fin nearly white in re- cently preserved specimens. In specimens preserved for over one year white may be nearly gone, leaving this area of fin dusky translucent. White on anterior portion of anal fin extending distally for more than half of length of fin rays, usually more than 75-80% of their length. Distal to white stripe, anal fin dusky hyaline except for in an occasional specimen having tips of fin rays and membranes between them more dusky than re- mainder of fin. Sometimes in males anterior 13-15 fin rays and their mem- branes distal to white stripe so densely covered with dark chromatophores as to appear black. Pectoral, pelvic, and caudal fins dusky hyaline. Dorsal fin with same color except for a large black spot covering two-thirds (in females) to three-fourths (in males) of distal distance of third to fifth or sixth fin rays and their interconnecting membranes. Distal half of second and, in females, third ray also white. Distal tip of fourth fin ray of females often white. These white areas translucent in specimens preserved for a great length of time. Formalin destroys white pigment in this species and in H. socolofi. Adipose fin dusky. Color in life——Brown to black pigment about same as described above under color in alcohol, but much silvery pigment in area of abdominal cavity, on head below eye, operculum, and proximal three-fourths of lower jaw. Both sexes appearing a “warm” but relatively pale chocolate brown color with an orange to pale red aspect on sides below junction of epaxial and hypaxial musculature. Above this junction, on back, color nearly same except orange and red colors more or less replaced by scattered pale green, blue, and gold chromatophores. Back considerably darker than remainder of body. A deep-red spot (often fully as brilliant as red in eye) just dorsal to or somewhat overlapping six (or often seventh) to ninth or tenth lateral- line scales. This spot may cover area of 1% to about 3 scales. A similar red color may occur at base of caudal fin but is very pale and often ob- scured by brown to black chromatophores. Just anterior to eye and posterior to dorsal one-third of premaxillary is a vertically elongate metallic gold or copper colored bar. Top of head dark greenish brown. Opercle slivery but often with a golden orange VOLUME 90, NUMBER 2 343 color and scattered brown to black chromatophores. Lower jaw, distal half of premaxillary, and ventral orbital region same color as opercle ex- cept distal tip of lower jaw greyish brown. Proximal half of premaxillary greyish dusky brown to black. Just posterior to dorsoposterior portion of eye occurs a reflectant golden-brown spot often obscured by dark brown pigment of head and back. Spot is slightly larger than pupil of eye. Fins dusky hyaline. Caudal, pectoral, and pelvic fins without other pigment. White of anal fin (distributed as described above under color in alcohol) with an intense milky appearance, especially on anterior 16 to 18 rays. In males white of stripe may be intense to posteriormost anal-fin ray. In females it is often intense only in region of third through eighth or twelfth rays. Scattered golden-orange chromatophores occur over re- maining portions of anal fin and are more dense on terminal 10 or 11 rays. Distal tips of anal-fin rays with membranes between rays nearly black. Adipose fin colorless except for dusky chromatophores. Dark spot of dorsal fin distributed as described above under color in alcohol. White areas proximal and distal to black spot may be pale milky orange or a deep rich milky orange in both sexes. Sometimes in males nearly as red as body spot. Sometimes females with a red stripe at base of dorsal fin covering both rays and interradial membranes, sometimes only rays. Eye same color as described for H. socolofi above. Discussion.—The two species of bleeding heart tetras described above can be distinguished by comparison with the diagnoses given above for each species. The new species, H. socolofi, usually has a proportionally shorter snout (6.9 to 7.9% of standard length compared to 7.8 to 8.5% in H. erythrostigma); shorter anterior dorsal-fin rays in adult males [30.4-34.4, x = 32.9 of SL, n = 3 (largest males) in H. socolofi compared to 38.3-50.5, * = 42.1% of SL, n = 4 (largest males) in H. erythrostigma]; shorter an- terior anal-fin rays in adult males [18.1-22.1, x = 20.4% of SL, n = 3 (largest males) in H. socolofi compared to 27.4-35.7, « = 30.9% of SL, n = 4 (largest males) in H. erythrostigma]; somewhat shorter pelvic-fin length in adult males (18.6—20.8, x = 19.7% of SL, n = 3 (largest males) of H. socolofi com- pared to 23.0-28.5, x = 24.7% of SL, n = 5 (largest males) of H. erythro- stigma]; numerous small bony hooks on the dorsal and anal fin of the males (nearly absent in the males of H. erythrostigma); and usually 32 vertebrae, one specimen with 33 (33 vertebrae in all specimens of H. erythro- stigma examined). Of the above characters the distribution of white pigment on the anal fin, the length of the anterior rays of the anal and dorsal tins in males (and to a much lesser extent in females) and the generous oc- currence of bony hooks in H. socolofi appear to offer the best differences between these two very closely related species. All the characters serving best to distinguish the species are associated with sexual dimorphism and ‘SG6T suLinp niog wo1y poytodun oq 0} pres yng uUMOUXUN APVOO] “Uotufoods wuMenbe “TS wu g'oPF ‘oeMloy ‘PFELT INNS ‘(49[A0,7 ) DUdYSOWYpita UOOhIgossaydhiyy °g “BI Z fe Fs © J rx WN Figs. 2-7. Male genitalia. Lepidostoma aztecum n. sp.: 2, Lateral; 3, Dorsal. L. heveli n. sp.: 4, Dorsal; 5, Lateral. L. steinhauseri n. sp.: 6, Dorsal; 7, Lateral. 379 VOLUME 90, NUMBER 2 380 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON and south to Guatemala as the records testify. There is some variation in the shape of the apical hooklike process of the clasper. In the series from Chiapas and Guatemala, and to a lesser degree, that from Veracruz, the apical half of this process in lateral aspect is sharply angled ventrad, but does not appear greatly different in ventral aspect. Considering that the other parts of the genitalia and the modifications of the antennae and maxillary palpi are the same in all this material, and that the modifications that do occur appear to be clinal in nature, we consider these to be all one species. Flint (1965, fig. 10) has provided figures of the male genitalia and Denning (1968, fig. 7) figures of the male basal antennal segment and female genitalia. Material—MEXICO, Chiapas, Dolores, route 190 km 1190 (20 km S of San Cristobal de las Casas), 8-9 July 1966, Flint & Ortiz, 1¢. Veracruz, La Joya, Rt. 140, km 307 (about 30 km NW of Jalapa), 21 July 1965, Flint & Ortiz, 86 22; same, but 1 July 1966, 26. GUATEMALA, Solola, 21 km’ Sof Nahwala, 9) Awee 1967, ©; 5, Flint, ji, 26. The species is also recorded from U.S.A., Arizona, and Mexico, Durango. Lepidostoma delongi Ross Lepidostoma delongi Ross, 1946, p. 283.—Fischer, 1970, p. 42. This species is known only by the unique male type taken at Zitacuaro, Michoacan, Mexico. The male genitalia were illustrated in the original descriptions (Ross, 1946, fig. 12). Lepidostoma frontale (Banks) Eremopsyche frontalis Banks, 1901, p. 367.—Fischer, 1970, p. 83. Lepidostoma frontalis: Flint, 1967B, p. 24. This species is a very close relative of L. mexicanum (Banks) as shown by the male genitalia (Flint, 1967B, fig. 124-125). The only significant differences between these species appear to be in the modifications of the male palpi, antennae and forewings. The basal segment of the antenna is much longer than the head, somewhat enlarged, and ventrally covered with long, scalelike setae that are directed anteriad forming a rather uni- formly smooth ventral surface, with the other surfaces more sparsely covered with setae projecting at all angles. The maxillary palpi are 1- segmented, spatulate, projecting anteriad, densely covered with elongate, widened setae directed dorsad and anteriad, the mesal faces are appressed, concave, and bear some short broad setae. The costal margin of the fore- wing is narrowly reflexed for its entire length, and the upper wing surface is covered with broadened hairs that are intensely purplish-iridescent at the proper angle. CN VOLUME 90, NUMBER 2 381 Material —MEXICO, Veracruz, La Joya, Rt. 140, km. 307 (about 30 km N.W. of Jalapa), 1 Aug. 1966, Flint & Ortiz, 1é. Described from Jalapa, Veracruz, September. Lepidostoma heveli Flint & Bueno, new species Figs. 4-5 This species, a member of the unicolor group, is very closely related to L. steinhauseri n. sp. and more distantly to L. aztecum n. sp. The position and shape of the basodorsal process and apical lobes of the clasper, and the lobes of the tenth tergum, are distinctly different in all three species. Adult.—Length of forewing, 11 mm. Color brown, antennae and legs stramineous; forewings brown. Maxillary palpus 1l-segmented, outer sur- face sparsely covered with erect hairs, basal % terete, apical third tri- anguloid with inner surface concave and bearing some short, broad seta; palpi not contiguous apically. Antenna with basal segment dark, slightly inflated, as long as head, with scattered erect, flattened setae. Forewing with costal cell narrowly reflexed for basal half, margin basally with a sparse fringe of long hairs, entire wing surface bearing erect spatulate hairs, a small area of small scalelike hairs on and adjacent to furcation of vein Rs and nearby on M. Male genitalia: Tenth tergum in lateral aspect di- vided into two pairs of processes, dorsalmost elongate, slender, narrowly divided middorsally, ventral process with an elongate, slender lobe from posterior margin and several points. Clasper with basodorsal process elongate, close to body of clasper, extending slightly beyond midlength of clasper, ventral arm of process appressed to clasper, bluntly pointed apically; apex with a small, fimbriate apicolateral lobe, a rounded, and com- pressed central lobe, and a rounded, depressed mesal lobe; with a sinuous basal connection to base of aedeagus. Aedeagus with an enlarged base, a central curved, tubular portion and dorsolateral swordlike processes. Material—Holotype male: GUATEMALA, Quiche, El Quiche, 7.3 km S$ Chichicastenango (14°54’N, 91°07’W), 2,400 mts., 28 May 1973, Erwin & Hevel. USNM Type 74074. Lepidostoma knulli Ross Lepidostoma knulli Ross, 1946, p. 280.—Flint, L967A, p. 175.—Fischer, ISAO), {Oo Bikes Lepidostoma leechi Denning, 1962, p. 37 [NEW SYNONYMY]. This species is closely related to L. bakeri Flint, not only on the basis of the general structure and appearance of the male genitalia, but also by the apparently identical modifications of the antennae and palpi. There 382 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON is some variation in the exact shape of the apicoventral margin of the tenth tergum, and in the length of the basodorsal arm and the apex of the clasper. However, these slight differences appear in all combina- tions, and the sexual modification of the palpi and antenna are the same throughout. We therefore synonymize leechi with knulli. The maxillary palpus of the male is l-segmented, spatulate, concave mesally and bearing a dense brush of long spatulate setae internally from the base. The basal segment of the male antenna bears mesobasally a short round projection and internally a small spherical structure that can be extruded through this projection. The male genitalia were well illustrated by Ross (1946, fig. 10), with additional figures of the male genitalia and basal antennal segment by Denning (1962, fig. 5) as leechi. Material—MEXICO, Michoacan, Parque Nacional Morelos, near Mo- relia, 14 July 1966, Flint & Ortiz, 16. Mexico, La Marquesa, Parque Nacional Las Cruces, 5-9 July 1965, Flint & Ortiz, 34; same but 13 July 1966, 346. The types of L. knulli were from U.S.A., Arizona, the types of L. leechi from Mexico, La Marquesa, and it has been recorded from Mexico, Durango. Lepidostoma lacinatum Flint Lepidostoma lacinatum Flint, 1967A, p. 175.—Denning, 1973, p. 142. This species appears to bear some relationship to L. rectangulare Flint, but is quite distinct in its combination of characteristics. The figures of Flint (1967A, fig. 24-25) and Denning (1973, fig. 12) illustrate the vari- ation in the male genitalia found to date. The maxillary palpi of the male are 1-segmented, with the apical *% tri- anguloid, concave mesally, and filled with short, broad, scalelike setae. The basal antennal segment is darkened, enlarged, and about as long as the head, the flagellar segments, especially basally are distinctly flattened. The species has been recorded from Mexico, Durango and Sinaloa, and U.S.A., Arizona. Lepidostoma mexicanum (Banks) Olemira mexicana Banks, 1901, p. 367. Atomyoides bispinosus Ulmer, 1911, p. 25.—Fischer, 1970, p. 70 [NEW SYNONYMY]. Lepidostoma mexicanum: Ross, 1946, p. 288.—Flint, 1967B, p. 24.— Fischer 11972; p. 59: Lepidostoma alexanderi Denning, 1962, p. 37.—Flint, 1967A, p. 175. Lepidostoma bispinosum: Denning, 1962, p. 39.—Flint, 1967A, p. 175. This species is closely related to L. frontale (Bks.), differing primarily in the secondary sexual characteristics of the male palpi, antennae and VOLUME 90, NUMBER 2 383 forewing. We have compared the males and females of this species throughout its rather extensive range and find no discernable pattern in the variations of the tenth tergite, nor does there appear to be any real variation in the female genitalia or secondary male sexual characters. Careful comparison of the female type of mexicanum with females as- sociated with males of the bispinosum form establishes the fact that these are the same species, and mexicanum being the oldest valid name it takes priority over the others. The maxillary palpi of the male are 1-segmented, with the apical half trianguloid, concave mesally, and filled with short, scalelike setae, the basal antennal segment is enlarged, as long as the head, and bears a covering of erect setae. The male of the “bispinosum” form was well illustrated by Ulmer (1911, fig. 12-13) and Denning (1962, fig. 7) with the “alexanderi” form shown by Denning (1962, fig. 4). The female genitalia were illustrated by Flint (1967B, fig. 126-127). Material—PANAMA, Chiriqui, E] Volcan, Chiriqui Viejo River, 5,280’ 22 July 1964, A. Broce, 16. Bambito, 1,500 m, 22 May 1973, G. Ekis, 14 12. Boquete, 16-17 July 1967, O. S. Flint, Jr., 36 22. COSTA RICA, Heredia, Vara Blanca, 24 June 1967, Flint & Spangler, 15¢ 42. Cartago, Navarro, July 1962, F. S. Blanton, 16. Volcan Irazu, 2,200-2,500 m, 27-28 May 1936, Reimoser, 2¢ (Vienna). GUATEMALA, Baja Verapaz, Purulha, Schaus & Barnes, 12. Izabal, Cayuga, Schaus & Barnes, 14. Quezaltenango, Volcan Santa Maria, Aug., Schaus & Barnes, 12. MEXICO, Hidalgo, 26.2 km N of Zimapan, 19 May 1973, Erwin & Hevel, 12. Michoacan, Parque Nacional Morelos, near Morelia, 14 July 1966, Flint & Ortiz, 3¢. The species has been recorded from Costa Rica, San Jose (type of bispinosa), Mexico, Distrito Federal, Tacubaya (type of mexicanum) and Durango, and U.S.A., Arizona (type of alexanderi). Lepidostoma rectangulare Flint Lepidostoma rectangulare Flint, 1967A, p. 176. This species appears to be most closely related to L. talamancense n. sp. from Costa Rica, but is distinguished by the shape of the apex of the clasper and tenth tergum. The male genitalia were illustrated by Flint (1967A, fig. 26-27). The species is still known only from the holotype from Durango, Mexico. Lepidostoma reimoseri Flint & Bueno, new species Figs. 10-11 This species, a member of the unicolor group, appears to be closely re- lated to L. rectangulare Flint and L. talamancense n. sp. From these, 384 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON reimoseri is easily recognized by the more elongate tenth tergum with setate papillae dorsally, and in the shape of the apical lobes of the clasper. Adult.—Length of forewing, 9 mm. Color brown, antennae stramineous; forewing covered with elongate, brown, scalelike hairs with small inter- spersed spots of stramineous scales, hind wing sparsely covered with brown hairs. Maxillary palpus 1l-segmented, very short, appressed to venter of head, inflated apicad, lateral and ventral faces covered with pale, clavate hairs. Antenna with basal segment terete, enlarged, as long as head, covered with dark, clavate hairs; flagellar segments with dark, clavate hairs dorsally. Fore and hind wings, dorsally covered with clavate hairs; foretibia and tarsus externally with some dark, clavate hairs. Male genitalia: Tenth tergum in lateral aspect 1% times as long as broad, rounded apically with setate papillae dorsad; in dorsal aspect deeply di- vided on midline, halves divergent. Clasper with basodorsal process semi- erect, extending to midlength of clasper, ventral arm of process appressed to clasper, rounded apically; apex with a small lateral flange, a small compressed central lobe, and a depressed mesal flap; with a straight basal connection to base of aedeagus. Aedeagus with an enlarged base, a central, curved tubular portion, and dorsolateral swordlike processes reach- ing to apex of tube. Material—Holotype, male: COSTA RICA, [Cartago], Volcan Irazu, 2,200-2,500 m, 21-28 May, 1930, Reimoser (Vienna Museum). Paratype: Same data, 1¢ (U.S.N.M.). Lepidostoma steinhauseri Flint & Bueno, new species Figs. 6-7 This species and L. heveli n. sp. are very closely related, and were at first believed to be the same species. There are differences between the two in the shape of the processes of the tenth tergum, the apical lobes of the clasper, and the height at which the basodorsal arm of the clasper is borne. However, more important to us than these differences which are only quantitative in nature, are the differences in the secondary sexual modifications. These differences are seen in the maxillary palpi, and especially the forewings which have a much more strongly reflexed costal cell, with a very long hair fringe, no short, scalelike setae, and a more re- stricted distribution of the spatulate hairs. Adult—Length of forewing, 12 mm. Color brown; antennae, legs, and thorax ventrally stramineous; forewing brown, mottled with stramineous hairs. Maxillary palpus l-segmented, shorter than head, outer surface covered with erect hairs; basal half terete, apical half trianguloid, with inner surface concave with small scalelike setae. Antenna with basal segment dark, slightly inflated, shorter than head, with long, erect setae. VOLUME 90, NUMBER 2 385 Figs. 8-11. Male genitalia. Lepidostoma talamancense n. sp.: 8, Lateral; 9, Dorsal. L. reimoseri n. sp.: 10, Lateral; 11, Dorsal. Forewing with costal cell narrowly reflexed for basal half and with mar- gin bearing a dense fringe of long setae; surface with many erect, spatulate hairs beneath fringe, and in a broad band from base almost to apex be- tween R3 and My, these hairs forming an enlarged hump at the base of M. Male genitalia: Tenth tergum in lateral aspect divided into two pairs of processes, dorsalmost elongate, divided middorsally, ventralmost with a narrow, lateral, winglike lobe and a short, terete ventral lobe. Clasper with basodorsal process elongate, raised above, but parallel to, body of clasper, 386 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON extending to midlength of clasper, ventral arm of process appressed to body of clasper, broadly pointed in dorsal aspect; apex with a large, fimbriate apicolateral lobe with a distinct basolateral lobe, an elongate, rounded and compressed central lobe, and a winglike depressed mesal lobe; with a sinuous basal connection to base of aedeagus. Aedeagus with an enlarged base, a central, curved, tubular portion, and broad, dorsolateral sword- like processes. Material—Holotype, male: EL SALVADOR, Santa Ana, Cerro Mira- mundo, 2,300 m, 23 Jan. 1971, S. Steinhauser. USNM Type 74075. Lepidostoma talamancense Flint & Bueno, new species Figs. 8-9 This species is closely related to L. reimoseri n. sp. and is probably its sister species found in the Cordillera de Talamanca. The primary dif- ferences between these two species lie in the tenth tergum which in talamancense is short and round apically, and in dorsal aspect has a dis- tinct mesal shoulder. The secondary sexual modification so apparent on the wings of reimoseri are lacking in talamancense. Adult.—Length of forewing, 9-13 mm. Color brown; antennae and legs stramineous; forewing either golden brown with dark flecks, or dark brown with scattered golden flecks, with a tuft of pale hairs on hind margin. Maxillary palpus l1-segmented, spatulate, outer surface covered with long, slightly flattened setae; inner face concave apically and filled with small, scalelike setae. Antenna with basal segment darkened, terete, slightly inflated, as long as head. Wings and legs without sexual modi- fications. Male genitalia: Tenth tergum in lateral aspect as high as long, apex rounded, serrate; in dorsal aspect with a distinct mesal angle. Clas- per with basodorsal process semierect, extending not quite to midlength of clasper, ventral arm of process appressed to clasper, broadly rounded apically; apex with a well-developed lateral lobe, a compressed central lobe, and a depressed mesal flap. Aedeagus with an enlarged base, a central, tubular, portion, and small dorsolateral process not reaching apex of central tube. Material—Holotype, male: COSTA RICA, Cartago, Ojo de Agua, route 2 km 75, 30 June 1967, Flint & Ortiz. USNM Type 7076. Paratype: Same data, 13 22; same, but 20 July 1967, 34 29. Literature Cited Banks, N. 1901. A list of neuropteroid insects from Mexico. Trans. Amer. Ent. Soe 27/-36l—o7 1s Cabrera, A. L., and A. Willink. 1973. Biogeografia de America Latina. Organization of American States, Washington, 120 pages. VOLUME 90, NUMBER 2 387 Denning, D. G. 1962. New Trichoptera. Pan-Pacific Ent. 38:33-39. 1964. Trichoptera of Baja California. Pan-Pacific Ent. 40:128-134. 1968. New species and notes of western Trichoptera. Jour. Kans. Ent. Soc. 41:63-69. 1973. New species of Trichoptera. Pan-Pacific Ent. 49:132-143. Fischer, F. C. J. 1970. Philanisidae, Lepidostomatidae, Brachycentridae, Beraeidae, Sericostomatidae, Thremmatidae, Helicopsychidae. Trichopt. Catalog. XI:1-316. Flint, O. S., Jr. 1965. New species of Trichoptera from the United States. Proc. Ent. Soc. Wash. 67:168—176. 1967A. Studies of Neotropical caddisflies, VI: On a collection from north- western Mexico. Proc. Ent. Soc. Wash. 69:162—176. 1967B. Studies of Neotropical caddisflies, V: Types of the species described by Banks and Hagen. Proc. U.S. Nat. Mus. 123( 3619 ):1-37. Ross, H. H. 1946. A review of the Nearctic Lepidostomatidae (Trichoptera). Ann. Ent. Soc. Amer. 39:265-291. 1951. The Trichoptera of Lower California. Proc. Calif. Acad. Sci. 27: 65-76. Schmid, F., and R. Guppy. 1952. An annotated list of Trichoptera collected on southern Vancouver Island. Proc. Ent. Soc. Brit. Columbia 48:41—42. Ulmer, G. 1911. Einige Siidamerikanische Trichopteren. Ann. Soc. Ent. Belgique 55:15-26. Smithsonian Institution, Washington, D.C. 20560 and Instituto de Bi- ologia, U.N.A.M., Apartado Postal 79-153, Mexico 20, D.F. PROC. BIOL. SOC. WASH. 90(2), pp. 388-399 MARINE CHLOROMONADS: MORE WIDELY DISTRIBUTED IN NERITIC ENVIRONMENTS THAN PREVIOUSLY THOUGHT Alfred R. Loeblich III, and Karen E. Fine In 1973 we examined a culture of a flagellate referred to as the “Long Island Sound dino” obtained from Luigi Provasoli. It was originally iso- lated and grown in DV medium (Provasoli and McLaughlin, 1963) by I. J. Pintner from a water sample collected by Robert Wassman at Larchmont, Long Island Sound, New York. At the time of collection it was abundant enough to discolor the water. Loeblich and Loeblich (1973) determined that this organism was Heterosigma akashiwo (Hada) Hada (1968), and concluded that it exhibited chrysophyte affinities. Loeblich (1976) sub- sequently considered this species to be a chloromonad and transferred it to the genus Chattonella Biecheler (1936) as Chattonella akashiwo (Hada) Loeblich (1976). Its possession of fucoxanthin, lack of condensed inter- phase chromosomes, and cell morphology different from the typical dino- flagellate motile cell suggested that it was not a dinoflagellate. The possession of multiple plastids appeared to exclude it from the Chrysophyta. We obtained a culture of another organism (strain FCRG 51, UTEX 2162) from James Jordan, who originally isolated it from 2 m depth off the sewage outfall at Pt. Loma, California, on July 10, 1970. Jordan had identi- fied it as a species of Exuviaella Tsenkovskii (1881). This organism may be the undescribed chloromonad reported by Lackey and Clendenning (1965) off Point Loma during their survey of the plankton of San Diego Bay, California. This isolate appeared from light microscopic observations not to be a dinoflagellate but instead to be related to Chattonella akashiwo. Electron and light microscopic observations indicated that strain FCRG 51 was identical to the marine chloromonad Fibrocapsa japonica Toriumi & Takano. In addition, Olisthodiscus luteus Carter strain “Olisth” was obtained from R. R. L. Guillard. It was originally isolated in 1952 by R. J. Conover from Long Island Sound at Milford, Connecticut. A survey of published literature suggests to us that enough similarities (many previously unrecognized) are present between members of the genera Chattonella |= Hornellia Subrahmanyan (1954); see Hollande and Enjumet (1956)], Fibrocapsa Toriumi & Takano (1973), Heterosigma Hada (1968) [invalid, International Code of Zoological Nomenclature, Art. 13 (b); International Code of Botanical Nomenclature, Art. 45], and Olisthodiscus Carter (1937) to allow these genera to be considered con- generic, representing several species of marine chloromonads. The purpose of this paper is to present electron and light microscopic VOLUME 90, NUMBER 2 389 observations that have facilitated the identification of strain FCRG 51 and to explain why we are transferring Fibrocapsa japonica, Heterosigma inlandica and Olisthodiscus luteus to Chattonella, a genus of marine chloromonads. All three isolates were grown in medium GPM (Loeblich, 1975) at 20°C under 225 ft-c cool white fluorescent light. Cells were fixed for electron microscopy by two methods. Method A is identical to the pro- cedure used for electron microscope sections of O. luteus by Leedale et al. (1970). Method B is given in Loeblich (1976). Observations were made using a Phillips 300 electron microscope. Description of Strain FCRG 51 The cells (Fig. 1) are anteriorly biflagellate and motile. They appear slightly flattened, and their mean length is 25.6 wm and their mean width (taken at the widest point) is 18.6 wm. The cells lack a cell wall or cover- ing of scales and possess only a cytoplasmic membrane (Fig. 4). Numer- ous plastids (mean number 29.4) surround a central nucleus, which has several golgi bodies capping the anterior nuclear surface (Figs. 2-4). The nucleus appears to have a small projection towards the flagellar region (Fig. 5). A similar feature was noted by Mignot (1976) in Chattonella subsalsa Biecheler (1936) and by Leadbeater (1969) in Olisthodiscus luteus. In both these species, the nucleus projects out at the site of attach- ment of the rhizoplast; the rhizoplast runs from the nucleus towards the flagellar bases. We were unable to find flagella intact in the sectioned material. It is highly likely, however, that the flagella are heterokont, as the cytoplasm contains numerous vesicles with microtubular contents ( Fig. 6) suggestive of stiff flagellar hairs. Stored flagellar hairs would be ex- pected for a heterokont. Similar vesicles occur only in those species bearing stiff flagellar hairs and have been detected in C. subsalsa by Mignot (1976) and in O. luteus by Leadbeater (1969) and Leedale et al. (1970); they are also common cytoplasmic components of freshwater chloromonads (Hey- wood, 1973). As further evidence of heterokonty, the flagella of strain FCRG 51 differ in function, one curving in an anterior direction, the other directed posteriorly or at a slight angle from the cell in a posterior direc- tion. The posteriorly directed flagellum does not appear to beat but instead seems to have a steering function since it is held rigid in varying angles from the cell surface during swimming. Similar observations have been made by Iwasaki (1971) for an organism identified as Exuviaella sp. which we believe on the basis of illustrations to be conspecific with isolate FCRG 51 and with F. japonica. The golden plastids are surrounded by four membranes; the outer two belonging to the plastid endoplasmic reticulum, the inner two representing PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 390 VOLUME 90, NUMBER 2 391 the organelle-bounding membranes (Fig. 7). Tubules are located between these two sets of membranes on the proximal or pyrenoid side of the plastids (Fig. 7). Gibbs (1962) found similar tubules in an identical loca- tion in O. luteus, strain “Olisth.” The two outer membranes are not in contact with the nuclear envelope as is the case for many heterokonts. Girdle lamellae are lacking or poorly developed in the plastid, and the pyrenoid is located on the proximal face of the plastid. An identical pyrenoid position was found in C. subsalsa by Mignot (1976) and in O. luteus by Gibbs (1962). The pyrenoid is traversed by lamellae composed of 2 or 3 thylakoids. Sections reveal a non-membranous, darkly staining band between the membranes bounding the proximal plastid face and the pyrenoid (Figs. 4, 7). This layer is present in all the plastids of this species, but is not evident in the published micrograph of the related species, O. luteus (Leadbeater, 1969) and C. subsalsa (Mignot, 1976). The storage product is not recognizable in the micrographs and is unknown. Bidwell (1957) has determined for the related species, O. luteus, that glucose is the main sugar in the insoluble carbohydrate fraction and d-mannitol the main soluble product of photosynthesis. When viewed in the light microscope, many refractile structures are present at the posterior end of the cell (Figs. 1-3). Upon cell lysis, these elongate considerably (up to 220 um). They are thought to be homologous to the ejectile mucocysts detected by Mignot (1976) in C. swbsalsa and to the mucilage vesicles that are discharged by O. luteus (Leadbeater, 1969). Distinct, mucocyst-like structures are not readily apparent in sectioned cells. Electron micrographs (Figs. 4, 6), however, do reveal a fibrous ma- terial abundant beneath the surface of the cell which may represent the refractile bodies evident in the light microscope (Figs. 1-3). These could be homologous to the fibrous mucocysts reported for other species. The length of the discharged mucocysts of strain FCRG 51 is greater than the 30 pm length of the discharged mucocysts of C. subsalsa (Mignot, 1976). Discussion of Marine Chloromonads Our study of strain FCRG 51, the “Long Island Sound dino,” and strain “Olisth,” and a review of the litearture lead us to several conclusions. < Figs. 1-4. Strain FCRG 51. Fig. 1, Motile cell with flagella at upper end. Arrow indicates mucocysts located in the posterior of cell. Scale line = 12.5 wm. Fig. 2, Par- tially squashed cell showing centrally located nucleus and 33 plastids. Arrow points to undischarged mucocysts. Scale line = 25 um. Fig. 3, Cell squashed to reveal dis- charged mucocysts (arrow). Scale line = 50 um. Fig. 4, Cell showing nucleus, cyto- plasmic membrane, and peripheral arrangement of the plastids. Three arrows indicate fibrous material, perhaps the mucocysts. Fixation by method A. White scale line = 5 um, PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 392 VOLUME 90, NUMBER 2 393 Strain FCRG 51 is conspecific with Fibrocapsa japonica. The organisms are neritic and share similar cell morphology, cell dimensions, location and type of nucleus, heterokont arrangement of flagella, and number and ar- rangement of plastids. Both lack a cell wall, and the location of mucocysts is the same in both, as is the manner of mucocyst discharge from the cell’s posterior. On the basis of morphology, the Exuwiaella species studied and illustrated by Iwasaki (1971, 1973) also appears to be conspecific with F. japonica. The illustrations presented by Iwasaki (1971) of another species, which he determined to be a euglenid of the genus Eutreptiella da Cunha (1914) look strikingly similar to those of C. swbsalsa given by Biecheler (1936). Strain FCRG 51 does not appear to be a dinoflagellate as thought by Jeffrey et al. (1975), but instead seems to be congeneric with C. subsalsa (= Hornellia marina Subrahmanyan, 1954), O. luteus, C. akashiwo, and Heterosigma inlandica. Each of these five species posseses two anteriorly inserted flagella that are heterokont in function and most likely in struc- ture (as proven for C. subsalsa and O. luteus). The species display a similar, somewhat elongate cell shape and lack a cell wall. Each contains a centrally located nucleus surrounded by multiple, golden plastids. Muco- cysts have been reported for C. subsalsa, F. japonica, and O. luteus. All of these species are neritic and at times extremely abundant (Iwasaki et al., 1968; Iwasaki and Sasada, 1969; Pratt, 1966). Several (e.g., C. swbsalsa and F’, japonica) have been associated with mortality of other marine animals (Subrahmanyan, 1954, Hollande and Enjumet, 1956, Iwasaki, 1971, Toriumi and Takano, 1973). Fucoxanthin has been detected in C. akashiwo by Loeblich and Loeblich (1973), in strain FCRG 51 by Jeffrey et al. (1975), and in O. luteus by Riley and Wilson (1967). Chlorophylls a and c have been detected by Jeffrey et al. (1975) for strain FCRG 51, and by Gibbs (1962), Ricketts (1966), Riley and Wilson (1967) and Jeffrey (1976) for O. luteus. Sufficient morphological similarities exist between Olisthodiscus luteus strain “Olisth,” Chattonella akashiwo (the Long Island Sound isolate), and the Japanese C. akashiwo that these organisms may represent the same species. Ample physiological information is available for the Japanese < Figs. 5, 6. Strain FCRG 51. Fig. 5, Nucleus containing a darkly stained nucleolus. Note the non-dinoflagellate features (lack of condensed, fibrous chromosomes). Upper right side of nucleus projects out towards the flagellar region. Fixation by method A. White scale line = 2.5 wm. Fig. 6, Cytoplasmic region where numerous vesicles occu with tubular contents (t). The tubules are thought to be stored flagellar hairs. Arrows point to fibrous bodies (mucocysts) similar to those described for other marine chloromonads. Fixation by method B. Scale line = 1 um. 394 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 7. Strain FCRG 51. Cross-section of plastids. Inner two membranes are the plastid envelope (pe); outer two are the plastid endoplasmic reticulum (per). Lamellae comprised of two thylakoids traverse the pyrenoid region (p). A non- membranous band (large arrows) occurs in the margin of the pyrenoid beneath the plastid envelope. Tubules (t) may occur between the plastid and the plastid envelope (pe). Fixation by method A. White scale line = 0.5 um. VOLUME 90, NUMBER 2 ‘ATOATOOdSoL soseyd YIAOIS IvouT] oye] pue Yro1s [euoucdxe BZurinp [joo 19d spyseyd Jo 1oqumu urout ay} Ske BET pure ['[Z spodas (OL6T) ‘TB 3 OoTOHRD ‘][90 19d spyseyd ¢Z—yL swoder (B9GT) AoOGpvaT ‘ZT ueY} o10W DABY S{[IO oB1e] SpOdeI (JEG) JopRy , ‘s][oo osevyd-soy oye] UO XOOF 7 poinsvout o19M (19 =U) Joquinu pysefd pue (gg =U) suOoIsUaUTIp [IAD , ‘GI ‘Bly Wo1y UoyRyL , (FO6L ) URAURLUYRIqnS sno.ouInu 9C-9T ce-OT €)-9G DULIDUL DYJAULO fF] (9€6T) t9poyporg sno1ounu — CGaGil 0S-OFE DSIDSGNS DIJAUOWDOY YD (LEGT) ToMRD »61—-G I=@ Il 6I-ZI snag pyauoyoyy C'9Z = ueIpou PST uerpour L'CZ = ueipow toded sty, P6¢=% = 981 =*% 9¢¢ = * aSV—LI a6 €C—-G'ET av GE—-9'6I IS DYWOH 9Fe[OST (CL6T) OuRyR], pure punto], = (lees LTRS 0-0 pawuodnl pyjauoywyy (S961) ®PPH 0S-O1 yy ZI-8 SI-O1 DIIPUD]UL DYJaU0JWwY (LO61T) BpeH «SI-FI = II-8 SI-ZI oMlyspyD D]]aU0WDYD a a eo ae ee SD ee ee JOUILIFOY roquimu SSOUYOTY Ypesig yjsue'T SUISIURBIC) Phseld (Wu UL) SUOISUSUIIp []eD a aa a ee ee ee a Bee ‘SPPUOWWOLOTYO OUMeUL JO SUOTILIOSap [VUISIIO oYyy pue TG OYOW Ulesjs Jo uostieduioo y “T aqGeL 396 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON isolate of C. akashiwo, while much morphological information is available for the English and American isolates of Olisthodiscus luteus. Until com- parable data are available we prefer to retain O. luteus and C. akashiwo as separate species. All of these flagellates most likely represent marine members of the division Chloromonadophyta. All exhibit features characteristic of both marine and freshwater chloromonads. The cells are flattened and motile, lack a cell wall, and can change their shape. At times they may exist as spherical, nonmotile cells. They are biflagellate and heterokont, with the flagella in a groove or slight gullet, the hairy flagellum directed anteriorly, and the smooth flagellum oriented posteriorly. Each cell possesses a nucleus capped anteriorly by golgi bodies. Multiple plastids are present, en- veloped by four membranes. No particulate storage products or eye spots have been detected. Finally, ejectile structures, or mucocysts, are common and display a species-specific arrangement. Previous workers have suggested a xanthophyte (Carter, 1937) or chryso- phyte (Gibbs, 1962) relationship for O. luteus. A relationship with these two algal divisions seems unlikely. The possession of multiple plastids, the presence of fucoxanthin, and the apparent absence of xanthophylls typical of a true chrysophyte (e.g. violaxanthin, antheraxanthin, neoxanthin, and zeaxanthin) or of a true xanthophyte (e.g. cryptoxanthin 5, 6 mono- and diepoxides, vaucheriaxanthin, diadinoxanthin, and heteroxanthin) are in- consistent with such affinities. (For a detailed survey of the chlorophylls and carotenoids of chloromonads, haptophytes, xanthophytes, eustigmato- phytes, chrysophytes and diatoms see Loeblich and Loeblich, 1977). A relationship between the Pyrrhophyta and Chattonella akashiwo and H. inlandica, as proposed by Hada (1967, 1968), also seems unsound. Chattonella akashiwo and H. inlandica appear closely related to C. sub- salsa, O. luteus, and F. japonica. These latter species share many non- dinoflagellate features. They possess plastids with envelopes formed by four membranes, contain stiff flagellar hairs, lack an amphiesma (the multi- membrane cell covering unique to dinoflagellates), and have nuclei whose chromosomes remain uncondensed during interphase. In addition, O. luteus and F. japonica possess fucoxanthin as a major photosynthetic pigment, and both the c; and cz forms of chlorophyll c (Jeffrey et al., 1975; Jeffrey, 1976). Photosynthetic dinoflagellates possess peridinin and only the cz form of chlorophyll c. Finally, the DNA of the “Long Island Sound” isolate of C. akashiwo has been characterized, as has that of two freshwater chloro- monads (Rae, 1976), Vacuolaria virescens Cienkowsky (1870) and Gonyo- stomum semen (Ehrenberg) Diesing (1866). No unsual bases were de- tected, as would be expected for dinoflagellates. No evidence is apparent from the electron microscopical observations of Mignot (1976) to support the suggestion of Margalef (1968) that Chat- VOLUME 90, NUMBER 2 397 tonella subsalsa is a colorless cryptomonad with several photosynthetic symbiotic “xanthelles.” Chattonella japonica, C. luteus and C. subsalsa lack the ultrastructural features of a cryptomonad (e.g. a cryptomonad type pellicle is absent), and at least two of these species possess the photo- synthetically active xanthophyll fucoxanthin (absent in cryptomonads ). On the basis of the above data we transfer the following three species into the genus Chattonella. Chattonella luteus (Carter) Loeblich & Fine, comb. nov. Basionym is Olisthodiscus luteus Carter, 1937, p. 19, pl. 3, figs. 25-40; pl. 8, figs. 19-24. Chattonella inlandica (Hada) Loeblich & Fine, comb. nov. Basionym is Heterosigma inlandica Hada, 1968, p. 4, text-figs. 5a—c. Chattonella japonica (Toriumi & Takano) Loeblich & Fine, comb. nov. Basionym is Fibrocapsa japonica Toriumi & Takano, 1973, p. 27, text- nese 2a=c, pls). 1, 2. Acknowledgments We thank R. R. L. Guillard, J. Jordan, and L. Provasoli for cultures, and M. F. Hedberg for technical assistance. Support by research grants from the National Science Foundation (BMS 74-22151) and the National In- stitute of General Medical Sciences, U.S. Public Health Service (GM 06637 ) is appreciated. Literature Cited Bidwell, R. G. S. 1957. Photosynthesis and metabolism of marine algae. I. Photo- synthesis of two marine flagellates compared with Chlorella. Can. Jour. Bot. 35:945-950. Biecheler, B. 1936. Sur une chloromonadine nouvelle d'eau saumatre Chattonella subsalsa n. gen., n. sp. Arch. Zool. Expér. Gén. 78:79-83. Carter, N. 1937. New or interesting algae from brackish water. Archiv Protistenk. 90:1-68, pls. 1-8. Cattolico, R. A., J. C. Boothroyd, and S. P. Gibbs. 1976. Synchronous growth and plastid replication in the naturally wall-less alga Olisthodiscus luteus. Plant Physiol. 57:497—503. Cienkowsky, L. 1870. Uber Pamellaceen und einige Flagellaten. Arch. Mikroskop. Anat. 6:421—438. da Cunha, A. M. 1914. Contribuicado para o conhecimento da fauna de Protozoairos do Brazil. II. Mem. Instituto Oswaldo Cruz 6:169-179. Diesing, C. M. 1866. Revision der Prothelminthen. Abtheilung: Mastigophoren. S. B. Akad. Wiss. Wien 52:287—401. Gibbs, S. P. 1962. Nuclear envelope-chloroplast relationships in algae. Jour. Cell Biol. 14:433-444. Hada, Y. 1967. Protozoan plankton of the Inland Sea, Setonaikai I. The Mastigo- phora. Bull. Suzugamine Women’s College, Nat. Sci. 13:1-26. —. 1968. Protozoan plankton of the Inland Sea, Setonaikai I]. The Mastigo- phora and Sarcodina. Bull. Suzugamine Women’s College, Nat. Sci. 14:1—28. 398 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Heywood, P. 1973. Intracisternal microtubules and flagellar hairs of Gonyostomum semen (Ehrenb.) Diesing. Br. Phycol. Jour. 8:43-46. Hollande, A., and M. Enjumet. 1956. Sur une invasion des eaux du Port d Alger par Chattonella subsalsa (= Hornellia marina Sub.) Biecheler. Remarques sur la toxicité de cette Chloromonadine. Stat. Aquiculture Péche Castiglione, New series : 273-280. Intemational Code of Botanical Nomenclature adopted by the eleventh International Botanical Congress Seattle, August 1969. 1972. pp. 1-426. International Code of Zoological Nomenclature adopted by the XV Intemational Con- gress of Zoology. 1961. pp. i-xvii, 1-176. Iwasaki, H. 1971. Studies on the red tide flagellates-VI. On Eutreptiella sp. and Exuviaella sp. appeared in Bingo-Nada, the Seto Inland Sea, in 1970 [in Japa- nese]. Jour. Oceanogr. Soc. Japan 27:152—157. 1973. The physiological characteristics of neritic red tide flagellates. Bull. Plankton Soc. Japan 19:104—114. , T. Fujiyama, and F. Yamashita. 1968. Studies on the red tide dinoflagellates I. On Entomosigma sp. appeared in coastal area of Fukuyama. Jour. Fac. Fish. Animal Husbandry, Hiroshima Univ. 7:259-267. , and K. Sasada. 1969. Studies on the red tide dinoflagellates-II. On Hetero- sigma inlandica appeared in Gokasho Bay, Shima Peninsula. Bull. Japanese Soc. Sci. Fish. 35:943—-947. Jeffrey, S. W. 1976. The occurrence of chlorophyll c: and cz in algae. Jour. Phycol. 12:349-354. , M. Sielicki, and F. T. Haxo. 1975. Chloroplast pigment patterns in dinoflagel- lates. Jour. Phycol. 11:374-384. Lackey, J. B., and K. A. Clendenning. 1965. Ecology of the microbiota of San Diego Bay, California. Trans. San Diego Soc. Nat. Hist. 14:9-40. Leadbeater, B. S.C. 1969. A fine structural study of Olisthodiscus luteus Carter. Br. Phycol. Jour. 4:3-17. Leedale, G. F., B. S. C. Leadbeater, and A. Massalski. 1970. The intracellular origin of flagellar hairs in the Chrysophyceae and Xanthophyceae. Jour. Cell Sci. 6: 701-719. Loeblich, A. R., III. 1975. A seawater medium for dinoflagellates and the nutrition of Cachonina niei. Jour. Phycol. 11:80—86. ——. 1976. Dinoflagellate evolution: speculation and evidence. Jour. Protozool. 23:13-28. , and L. A. Loeblich. 1977. Divisions Bacillariophyta, Chloromonadophyta, Chrysophyta, Eustigmatophyta, Haptophyta, Xanthophyta. CRC Handbook of Microbiology, 2nd ed., ed. by A. I. Laskin and H. A. Lechevalier, CRC Press Inc., Cleveland, Ohio: in press. Loeblich, L. A., and A. R. Loeblich III. 1973. A search for binucleate or chrysophyte containing dinoflagellates. Jour. Protozool. 20:518. Margalef, R. 1968. Présence de Chattonella subsalsa Biecheler dans le port de Barcelone. Rapp. Comm. Int. Mer. Médit. 19:581-582. Mignot, J.-P. 1976. Complements a l’étude des chloromonadines ultrastructure de Chattonella subsala Biecheler flagellé d’eau saumatre. Protistologica 12:279-293. Pratt, D. M. 1966. Competition between Skeletonema costatum and Olisthodiscus luteus in Narragansett Bay and in culture. Limnol. Oceanogr. 11:447-455. Provasoli, L., and J. J. A. McLaughlin. 1963. Chapter 10. Limited heterotrophy of some photosynthetic dinoflagellates. In: Symposium on Marine Microbiology, ed. by C. H. Oppenheimer, pp. 105-113, C. C. Thomas, Springfield, II. VOLUME 90, NUMBER 2 399 Rae, P. M. M. 1976. Hydroxymethyluracil in eukaryote DNA: a natural feature of the Pyrrophyta (Dinoflagellates). Science 194:1062-1064. Ricketts, T. R. 1966. On the chemical composition of some unicellular algae. Phyto- chem. 5:67-—76. Riley, J. P., and T. R. S. Wilson. 1967. The pigments of some marine phytoplankton species. Jour. Mar. Biol. Ass. U.K. 47:351-362. Subrahmanyan, R. 1954. On the life-history and ecology of Hornellia marina gen. et sp. nov., (Chloromonadineae), causing green discoloration of the sea and mortality among marine organisms of the Malabar Coast. Indian Jour. Fish. 1: 182-203. Toriumi, S., and H. Takano. 1973. Fibrocapsa, a new genus in Chloromonadophyceae from Atsumi Bay, Japan. Bull. Tokai Regional Fisheries Res. Lab. 76:25-35. Tsenkovskiy, L. [Cienkowsky, L.]. 1881. Otchet” o byelomorskoy ekskursii 1880 g. Trudy Sankt-Peterburgskago Obshchestva Estestvoispytateley. 12:130-171, pls. 1-3. The Biological Laboratories, Harvard University, Cambridge, Massachu- setts 02138. PROC. BIOL. SOC. WASH. 90(2), pp. 400-411 A REDESCRIPTION OF OITHONA DISSIMILIS LINDBERG 1940 WITH A COMPARISON TO OITHONA HEBES GIESBRECHT 1891 (CRUSTACEA: COPEPODA: CYCLOPOIDA) Frank D. Ferrari Surface zooplankton samples were collected between 1400 & 1900 h on 9 & 18 April 1974 in Pago Pago Harbor, Tutuila Island, American Samoa. Pago Pago Harbor is shaped like an inverted “L”. The upright column of the “L” comprises the harbor mouth on the southern coast and is 1.5 km wide at this point. The column runs north 3 km where it is joined on its west side by the narrower, shallower base of the “L,” about 1 km long. In addition to detrital laden runoff entering the harbor from several streams, a tuna processing plant on the north shore, toward the head of the har- bor, disposes wastes into this area. These consist of blood, oil, and entrails of the processed fish. Oithona dissimilis Lindberg 1940 was the most abun- dant animal in the samples taken near the surface at the head of the harbor with an open conical net (mesh width 60 wm). Adult males and females with egg sacs, as well as many copepodid stages were present. Very few specimens were present in samples from the mouth of the har- bor and outside of the harbor. Oithona dissimilis Lindberg 1940 Figs. 1-3B Oithona dissimilis Lindberg, 1940:520, fig. 2; 1950:274 (key); 1955:467 (key). Oithona dissimilis oceanica Lindberg, 1947:52, fig. 2. Oithona hebes Giesbrecht 1891.—Wellershaus, 1969:276, figs. 88-102. Diagnosis.—Female length 0.61-0.70 mm (based on 30 specimens); Pr 1.3x Ur; cephalosome distinctly constricted dorsally and truncated an- teriorly; laterally attaining a small acute point which extends ventrally between first antennae. Greatest width at posterior end of cephalosome; width of metasome segments decreasing posteriorly. Metasome segments 1, 3, 4 each with a pair of dorsal sensory hairs, segment 2 with 2 pairs. As seen dorsally, with one group of sensory hairs in the middle of Url and a second group toward the posterior margin. Caudal ramus (Fig. 2A) 3X as long as wide, equal in length to anal segment. Al reaching posterior mar- => Fig. 1. Oithona dissimilis. A-—C, Female: A, Lateral; B, Dorsal; C, Ur, lateral. D-F, Male: D, Dorsal; E, Lateral; F, Cephalosome, ventrolateral; F, Ur, ventral. VOLUME 90, NUMBER 2 40] 402 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON gin of metasome segment 3. Bspd2 of Md (Fig. 2B) with 2 similar, slender curved spines armed with denticles; Ri with 5 setae, ultimate with setules. Large spines of Mx2 (Fig. 3A) with notch on dorsal margin and small denticles on distal fourth of ventral margin. Excluding terminal spine, Re P1-4 with 1-1-3, 1-1-3, 1-1-3, 1-1-2 external spines and 1-1-4, 1-1-5, 1-1-5, 1-1-5 internal setae; Ri P1l-4 with 0-0-1, 0-0-1, 0-0-1, 0-0-1 external setae and 1-1-5, 1-2-5, 1-2-5, 1-2-4 internal setae. External spines and terminal spine of Re P1 distinctly attenuated beyond hyaline membrane. Three internal setae of Ri P4 modified (Fig. 3B); proximal seta of Ri2 with tiny mem- branous flange on medial edge of tip; distal seta slightly curved with larger membranous flange; proximal seta of Ri3 P4 toward its distal end strongly curved, with a very large membranous flange medially. These setae are poorly developed in copepodid V. Internal setae of Rel P1-4 reduced. P5 with 2 setae, one each on a small lobe dorsally and on larger lobe ventrally. Caudal ramus, external apical seta %x as long as internal apical seta, equal in length to external seta. Genital opening with 2 spines, ventral one larger; a slight prominence ventral to area of opening. Male length 0.64-0.72 mm (based on 30 specimens); Pr 1.3 Ur; cephalo- some, in dorsal view distinctly constricted anteriorly, relatively wider than female; laterally anterior margin of cephalosome rounded, not as angular as female. Greatest width at posterior edge of cephalosome; width of first 2 metasome segments subequal; width of last 2 decreasing. Pr dorsally with slightly different arrangement of sensory hairs (Fig. 1D). Pr laterally (Fig. 1F) with a very complex group of integumental organs in an area comprising posteroventral part of cephalosome and posterior extension or flap of cephalosome overlapping following segment. These organs ap- parently composed of thickened base and thinner long hair, the latter usually broken. They adom posterior and ventral edges of flap and con- tinue anteriorly along ventral edge of cephalosome for half its length; dorsally along this length are 12 vertical rows; dorsal to these rows on flap 2 transverse rows, and on cephalosome a more or less longitudinal row, continuing anteriorly as a more curved, double row. Length of caudal ramus (Fig. 2D) 2x width, equal to length of anal segment. Bspd2 of Md with 2 curved spines, relatively shorter than female, armed with thinner denticles; Ri with 5 setae. Re and Ri P1-4 armed as female; all internal setae of Ri P4 unmodified. P5 with 2 setae laterally; genital flap with 1 small naked seta and ventrally a larger setuled one. Dorsal seta of caudal ramus reduced, external apical seta longer than external seta. > Fig. 2. Oithona dissimilis. A-B, Female: A, Caudal ramus, ventral; B, Md. C-E, Male: C, Md; D, Caudal ramus, ventral; E, Al. 403 NUMBER 2 VOLUME 90, 404 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Discussion Lindberg (1940) illustrated and briefly described Oithona dissimilis from the nearshore waters of Pondichery, India. In the description he overlooked an external spine on Re3 P1, but corrected this omission when he (1947) provided a more complete description of a subspecies, O. dis- similis oceanica, from Ratnaguiri, India. Lindberg remarked that the basis for the establishment of this subspecies was a difference in habitat. O. dissimilis was collected from the muddy brackish waters of the litoral zone of lagoons whereas O. dissimilis oceanica was found in the pelagic waters of the open sea. Lindberg had at his disposal only 3 females and 2 males of O. dissimilis oceanica. Based on records of specimens at my disposal and the co-occurrence of males, gravid females and copepodids, I concur with Lindberg’s original observation that O. dissimilis is an inhabitant of the brackish, muddy waters of bays and estuaries probably throughout the tropical Indo-Pacific region. Thus, the few representatives of the sub- species were probably members of a local inshore population swept out into the neritic zone. The apparent habitat difference alone should not be used as a basis for establishing this subspecies. Except for the mistake in the number of external spines of Re P1, Lind- berg’s (1940) discussion and illustrations fit the Pago Pago Harbor speci- mens. By a lapsus Lindberg labeled Re P4 as Ri P4. Differences do occur in Lindberg’s (1947) later more careful observations. There he stated that O. dissimilis oceanica lacks a seta on Rel P1, possesses a short, strong seta on Rel P2, a rudimentary one on Rel P3 and a more developed one on Rel P4; only P2 is illustrated. The genital flap of the male is illustrated with only 1 large seta and ventrally 3 small points. Although I have been unable to obtain specimens from Pondichery or Ratnaguiri, I have assumed a continuous distribution throughout the nearshore coastal area of India for O. dissimilis. I have had the opportunity of examining a collection of plankton samples from the Cochin Backwater of India taken by Dr. Thomas E. Bowman on 22 November 1968. In a sample collected in water of 27.3%c, a number of specimens of O. dissimilis were found which agree with those from Pago Pago Harbor, including Ri P4, genital opening of the female, and Al and genital flap of the male. The development of the flap on the cephalosome and the general arrangement of the integumental organs in the male are identical. I feel that the discrepancies between the absence or development of setae on Rel P1-4 and the male genital flap are too slight to warrant separation of the populations from Pago Pago Harbor and => Fig. 3. A-B, Oithona dissimilis, Female: A, Mx2; B, P4, anterior. C-—D, Oithona hebes, Female: C, Mx2; D, P4, anterior. —=~’ SS ’ ==5 a eS Les NN — ot | SSS SEE ZZ Sey ——— Fi = hs = - =< ee eS) Ny ‘ AN SS rs ey eS Ly — Se, 406 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Cochin from the populations described by Lindberg. Of these characters | only the morphology of the genital flap has any systematic value and this | difference could represent simply an error in observation. | Wellershaus (1969) described and illustrated as Oithona hebes Gies- brecht 1891, a species from Cochin Backwater, India. There are obviously awkward zoogeographic complications in finding a species originally de- scribed from the Bay of Guayaquil, Equador, and probably indigenous to the western tropical coast of the Americas, across the East Pacific Barrier in the Indo-Pacific region. Dr. Bowman has also made available to me plankton samples from the Bay of Guayaquil, Ecuador, collected in Feb- ruary, 1963. In these plankton samples from the type-locality of O. hebes, I have found numerous females and eight males of this species which agree with the descriptions of Giesbrecht (1892) and Kiefer (1936). Gonzalez and Bowman (1965) have briefly discussed differences between O. hebes specimens from the type-locality and Lindberg’s (1940) description of O. dissimilis and their observations are included in the following description. Oithona hebes Giesbrecht 1891 Figs. 3C-5 Oithona hebes Giesbrecht 1891:475; 1892:538, 549, pl. 34, figs. 8-9.— Grandori, 1912:15.—Farran, 1913:191 (key ).—Rosendorn, 1917:44.— Pesta, 1921:551, fig. G5.—Kiefer, 1929:9; 1936:320, figs. 1-5.—Rose, 1933:280, fig. 354—Lindberg, 1950:274 (key); 1955:467, (key).—Loffler, 1963:209.—Gonzalez & Bowman, 1965:269, fig. 19a-l, fig. 20a—b.— Bacon, 1971:85.—Santos, 1973:430, 432. Oithona minuta T. Scott 1894.—Coker & Gonzalez, 1960:8. (not) Oithona hebes Giesbrecht 1891.—Wilson, 1942:196. (not) Oithona hebes Giesbrecht 1891.—Wellershaus, 1969:276, figs. 88-102. Diagnosis.—Female 0.49-0.58 mm (based on 30 specimens). Pr 1.5x Ur; dorsally, cephalosome pointed anteriorly; dorsally, Url with 6-8 sensory hairs in 2 rows along midline. Ri of Md (Fig. 5B) with 4 setae, ultimate with setules; Bspd2 with 2 thick spines armed with numerous fine denticles. Mx2 (Fig. 3C) without notch or denticles on large spines. Ri P4 (Fig. 3D) with 2 modified setae; distal seta of Ri2 thicker and more strongly curved than proximal seta of Ri3 which is thinner and only slightly curved; both have membranous flange medially. Genital opening with a small point. Male 0.47-0.50 mm (based on 8 specimens). Integumental organs > Fig. 4. Oithona hebes. A-B, Female: A, Dorsal; B, Ur, lateral. C-—F, Male: C, Dorsal; D, Cephalosome, ventrolateral; E, Ur, lateral; F, Al. VOLUME 90, NUMBER 2 407 408 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON of cephalosome (Fig. 4D) in dorsal longitudinal row doubled only toward anterior end; ventral to this 11 vertical rows and 1 oblique row between third and fourth vertical rows. Ri of Md (Fig. 5D) with 4 setae; distal 2 spines of Bspd reduced, only one armed with denticles. Genital flap with 1 large seta with setules and smaller seta ventrally. Recently, Dr. Stefan Wellershaus of the Institut fir Meeresforschung, Bremerhaven, kindly sent me an aliquot of a sample collected on 31 May 1966 from the Cochin Backwater. From this I have removed and identified males and females of O. dissimilis which agree in all respects with those populations described above from Cochin and Pago Pago Harbor. Comments I have notes on five species of Oithona s.s. with modified setae on Ri P4. There are then the proximal inner seta on Ri3 P4 in O. brevicornis Giesbrecht (1891), noted and illustrated by Wellershaus (1969), the distal seta of Ri2 and proximal seta of Ri3 P4 in O. hebes Giesbrecht (1891) and O. colcarva Bowman (1975), and both setae of Ri2 and the proximal seta of Ri3 P4 in O. dissimilis Lindberg (1940) and O. plumifera Baird (1843), the latter noted and illustrated by Giesbrecht (1892). In each case, the number of setae modified and the extent of modification are specific for the species. The significance of these modified setae may be explained in this way. In the most diverse of the free-swimming copepods, the calanoids, many species exhibit various modifications of both male fifth legs and one of the first antennae. These appendages function to grasp and hold the female and help position the male during copulation. The spermatophore is probably transferred by the left fifth leg which has modified spines or, more often, ciliated parts of the distal segments of the limb to facilitate the handling of this delicate structure (for example, see fig. 22 of Vervoort, 1963). Lee (1972) has given a more complete discussion of the appendage modifications and inferred copulatory behavior of the Centropagidae. In the males of Oithona the first antennae are symmetrical and di- geniculate. A proximal series of segments can be rotated up to 270° in some species while the ultimate and pentultimate together can be folded 180° back onto the 3 segments proximal to them. Except for these append- ages, located far from the genital segment, the males do not possess other appendages modified to hold and transfer the spermatophore. It is possible that the female takes a much more active role in copulatory behavior, using her fourth legs and their modified setae to help transfer the spermatophore. > Fig. 5. Oithona hebes. A-B, Female: A, Caudal ramus, ventral; B, Md. C—D, Male: C, Caudal ramus, dorsal; D, Md. VOLUME 90, NUMBER 2 409 Ss > mm eS 410 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fleminger (1973) presented a general discussion of integumental organs in calanoid copepods and demonstrated the systematic value of the pore signature patterns for the genus Eucalanus. As yet I have been unable to describe the exact morphology of these organs in the Oithona males using a light microscope. However, I hope that use of a scanning electron micro- scope will clarify many points. I have examined males of O. plumifera, O. nana, and O. simplex from American Samoa. These animals exhibit nu- merous interspecific differences not only in the pattern and number of integumental organs, but also in the degree of development of the flap on the cephalosome. Fleminger (1973 & 1975) and Fleminger & Hulsemann (1973) have pointed the importance of prezygotic mating barriers in the selection process for several free-living calanoid genera. If this concept can be extended to Oithona, then the systematic value of the modified setae of the fourth legs of the female and the first antenna and integumental organs of the male becomes more apparent. Literature Cited Bacon, P. 1971. Plankton studies in a Caribbean estuarine environment. Caribbean Journal of Science 11 (1-2) :81-89. Bowman, T. 1975. Oithona colcarva, n. sp., an American copepod incorrectly known as O. brevicornis (Cyclopoida:Oithonidae). Chesapeake Science 16( 1) :134—137. Coker, R., and J. Gonzalez. 1960. Limnetic copepod populations of Bahia Fosfores- cente and adjacent waters, Puerto Rico. Journal of the Elisha Mitchell Scientific Society 76( 1) :8—28. Farran, G. 1913. Plankton from Christmas Island, Indian ocean, 2: On Copepoda of the genera Oithona and Paroithona. Proceedings of the Zoological Society of London 1913:181-193. Fleminger, A. 1973. Pattern, number, variability, and taxonomic significance of integumental organs (sensilla and glandular pores) in the genus Eucalanus (Copepoda, Calanoida). Fishery Bulletin 71(4) :965—1010. —. 1975. Geographical distribution and morphological divergence in American coastal-zone planktonic copepods of the genus Labidocera. Estuarine Research 1:392-419. , and K. Hulsemann. 1975. Systematics and distribution of the four sibling species comprising the genus Pontellina Dana (Copepoda, Calanoida). Fishery Bulletin 72( 1) :63—120. Giesbrecht, W. 1891. Elenco dei Copepodi pelagici raccolti dal tenente de vascello Gaetano Chierchia durante il viaggio della R. Corvetta “Vettor Pisani” negli anni 1882-1885, e dal tenente di vascello Francesco Orsini nel Mar Rosso, nel 1884. Atti Rendiconti della Rome Accademia dei Lincei, ser. 4, vol. 7, sem. 1, pp. 474-481. —. 1892. Systematik und Faunistik der pelagischen Copepoden des Golfes von Neapel und der angrenzenden Meersabschnitte. Fauna und Flora des Golfes von Neapel, monograph. 19, 831 pp., 541 pl. Gonzalez, J., and T. Bowman. 1965. Planktonic copepods from Bahia Fosforescente, VOLUME 90, NUMBER 2 4\] Puerto Rico, and adjacent waters. Proceedings of the United States National Museum 117( 3513) :241-304. Grandori, R. 1912. I copepodi. Pp. 1-41 in Carazzi and Grandori. Ricerche sul plancton della Laguna Veneta, vii + 64 pp. Kiefer, F. 1929. Copepoda, 2:Cyclopoida Gnathostoma. Vol. 53 in Das Tierreich, xvi + 102 pp. ——. 1936. Brasilianische Ruderflusskrebse (Crustacea Copepoda), gesammelt von Herr Dr. Otto Schubert. 3: Zur Kenntnis der Oithona hebes Giesbrecht. Zoologisher Anzeiger 114 (11-12) :320-322. Lee, C. 1972. Structure and function of the spermatophore and its coupling device in the Centropagidae (Copepoda:Calanoida). Bulletin of Marine Ecology 8: 20: Lindberg, K. 1940. Cyclopoides (Crustacea Copépodes) de Inde. Records of the Indian Museum 42(3):519-526. —., 1947. Cyclopoides (Crustacea Copépodes) nouveaux de lInde. Records of the Indian Museum 45(1):47—55. ——., 1950. Cyclopoides nouveaux ou peu connus (Crustacés Copépodes). Mém- oires Museum National d’Histoire Paris, new series, 29(3) :259-297. ——. 1955. Cyclopoides (Crustacés Copépodes) du Mexique. Arkiv for Zoologi, series 2, 7(23):459-489. Loffler, H. 1963. Zur Ostrakoden und Copepodenfauna Ekuadors. Archiv fiir Hydro- biologie 59(2) : 196-234. Pesta, O. 1920. Die Planctoncopepoden der Adria. Zoologische Jahrbiicher, System- atik 43:471-660. Rose, M. 1933. Copépodes pélagiques. Fauna de France, no. 26, 374 pp. Rosendom, I. 1917. Die Gattung Oithona. Wissenschaftliche Ergebnisse der Deut- schen Tiefsee-Expedition 23:1—58. Santos, J. 1973. Estudo preliminar, principalmente do plancton, das Aguas da Baia de Todos os Santos. Boletim de Zoologia e Biologia Marinha (Nova Serie) 30: 419-447. Vervoort, W. 1963. Pelagic Copepoda. Part I. Copepoda Calanoida of the families Calanidae up to and including Euchaetidae. Atlantide Report 7:77—194. Wellershaus, S. 1969. On the taxonomy of planctonic Copepoda in the Cochin Back- water (a South Indian Estuary). Ver6ffentlichungen Institute fur Meeres- forschung Bremerhaven 11:245—286. Wilson, C. 1942. The copepods of the plankton gathered during the last cruise of the Carnegie. No. 1 of Biology in Scientific results of Cruise 7 of the Carnegie during 1928-1929 under the command of Capt. J. P. Ault (Carnegie Institute of Washington Publication 536), v + 237 pp. Smithsonian Oceanographic Sorting Center, Smithsonian Institution, Washington, D.C. 20560. PROC. BIOL. SOC. WASH. 90(2), pp. 412-419 A NEW CRAYFISH (DECAPODA: CAMBARIDAE) FROM SAN LUIS POTOSI, MEXICO Horton H. Hobbs, Jr. The occurrence of the crayfish described herein in San Luis Potosi repre- sents an extension of the southern limit of the range of the Subgenus Scapulicambarus (those members of the genus Procambarus in which the male possesses hooks on the ischia of the third and fourth pereiopods and the first pleopod bears a distinct shoulder on the cephalic surface some distance proximal to the terminal elements). Only one other member of the subgenus, Procambarus (S.) clarkii (Girard, 1852), has been reported from Mexico. Hobbs (1962:273, 275) recorded its presence in the states of Chihuahua and Sonora. Specimens of the new species were obtained by Ned E. Strenth of the Marine Biomedical Institute, Galveston, Texas, and Thomas G. Littleton, of the University of Texas Health Science Center, San Antonio, from a small stream in the Panuco River basin, 15 miles (24 kilometers) west of Ciudad Valles. I wish to thank them for donating these crayfish to the Smithsonian and for permitting me to describe the species. Appreciation is also extended to Fenner A. Chace, Jr., and Isabel Pérez Farfante for their criticisms of the manuscript. Procambarus (Scapulicambarus) strenthi, new species Figs. 1-2 Diagnosis —Body pigmented, eyes well developed. Rostrum with small marginal spines at base of short, broad, triangular acumen, median carina absent. Carapace usually with cervical spine or tubercle (multiple spines in one specimen). Areola 6.9 to 9.2 times as long as broad and constituting 33.2-35.5% of total length of carapace (41.6-44.6% of postorbital carapace length). Suborbital angle weak and rounded. Postorbital ridge with cephalic spine or tubercle. Hepatic area with tubercles; branchiostegal spine present. Antennal scale almost twice as long as broad, widest at mid- length. Ischia of third and fourth pereiopods of first form male with simple hooks overreaching basioischial articulation, hook on neither ap- pendage opposed by tubercle on corresponding basis; coxa of fourth pereiopod with very prominent but thin boss on caudomesial ventral angle. First pleopods of first form male reaching coxa of third pereiopod, strongly asymmetrical, bearing prominent angular shoulder on cephalic surface, and provided with subapical setae (plumose ones mesially and stiff simple ones laterally) largely obscuring terminal elements; latter consisting of strong acute mesial process directed distolaterally, broad delicate lamel- late cephalic process with corneous margin partly surrounding large, VOLUME 90, NUMBER 2 413 distally truncate, corneous, dentiform central projection, and smaller comeous, subtriangular caudal process arising from rudimentary caudal knob on caudolateral extremity of pleopod. Annulus ventralis almost 3 times as broad as long, almost symmetrical in outline, with median area ele- vated and bearing sinuous sinus, latter originating in oblique groove extend- ing caudodextrally from cephalic median margin and terminating be- fore reaching caudal margin of annulus; tongue broad and short. First pleopod present in female. Holotypic male, form I.—Cephalothorax (Fig. la, g) subcylindrical, greatest width little more than height at caudodorsal margin of cervical groove. Abdomen slightly narrower than thorax (10.9 and 11.7 mm). Areola 9.2 times as long as wide with 1 or 2 widely spaced punctations across narrowest part. Cephalic section of carapace almost 3 times as long as areola, latter comprising 34.2% of entire length of carapace (43.5% of postorbital carapace length). Surface of carapace conspicuously punctate dorsally and tuberculate laterally except in orbital region where punctate. Rostrum broad basally, with weakly convergent margins bearing small marginal spines at level of midlength of penultimate podomere of an- tennular peduncle, acumen reaching midlength of ultimate podomere; mar- gins not thickened and not conspicuously elevated; upper surface concave, sparsely marked by small punctations anteriorly and larger ones basally. Subrostral ridge very weak and scarcely evident in dorsal view. Postorbital ridge moderately well developed, grooved dorsolaterally, and terminating cephalically in small spine. Suborbital angle weak and rounded. Branchio- stegal spine moderately well developed. Cervical spine represented by tubercle. Abdomen longer than carapace (26.8 and 24.3 mm). Pleura of third through fifth abdominal segments subtruncate ventrally and rounded caudoventrally. Cephalic section of telson with 3 spines (mesialmost on right vestigial) in each caudolateral corner, lateral and mesial spines im- movable. Cephalic lobe of epistome (Fig. lc) broadly subtriangular with cephalomedian projection; median part somewhat convex (ventrally) and bearing widely spaced setae; median fovea on main body of epistome dis- tinct and rather deep; epistomal zygoma broadly arched. Mesioventral surface of proximal podomere of antennular peduncle with strong spine at about midlength. Antennal peduncle with spines on basis and ischium; flagellum broken, but in paratypic male reaching caudally to fourth ab- dominal tergum. Antennal scale (Fig. 1f) 1.9 times as long as broad, widest at midlength; greatest width of lamellar area about 1.7 times that of thickened lateral part. Third maxilliped extending to level of distal end of proximal podo- mere of antennular peduncle; ischium with distolateral extremity angular 414 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON VOLUME 90, NUMBER 2 415 but not produced, its lateral half bearing scattered tufts of short simple setae; mesial half with clusters of stiff setae. Right chela (Fig. 1h) subovate in cross section, not strongly compressed. Entire palm with squamous tubercles, mesial surface with several irregular rows, mesialmost consisting of 8. Both fingers with low longitudinal ridge (that on dactyl somewhat indistinct) flanked proximally by tubercles and by punctations along most of length. Opposable margin of fixed finger with row of 8 (7 on left) tubercles (second and third from base larger ) along proximal half, large tubercle situated more ventrally slightly distal to midlength, and broad band of minute denticles present along almost entire length of finger. Opposable margin of dactyl with 2 rows of tubercles: dorsal row of 10 extending along basal half, and ventral row of 3 (left with only 2) along proximal fourth of finger; very broad band of minute den- ticles extending along distal four-fifths of margin. Carpus of cheliped longer than broad, lacking usual oblique furrow on dorsal surface; mesial surface with only 1 tubercle conspicuously larger than others nearby, and subacute tubercle present on dorsomesial distal angle; ventrodistal surface with usual 2 tubercles; remainder of podomere with setiferous punctations. Merus tuberculate dorsally, distomesially, and ventrally; 1 dorsal tubercle large and more spiniform than others on dorsodistal surface; ventral sur- face with irregular mesial row of 14 tubercles, lateral one of 11, and 2 ar- ranged obliquely between distal members of rows (left with 16, 12, and 3, respectively ), also additional tubercles flanking rows, and spiniform tuber- cle on ventrolateral extremity. Ischium with row of 4 (3 on left) tubercles ventromesially. Hooks on ischia of third and fourth pereiopods (Fig. le) simple, both overreaching basioischial articulation, neither opposed by tubercle on corresponding basis. Coxa of fourth pereiopod with moderately strong, vertically disposed caudomesial boss. Coxa of fifth with flattened (in longitudinal axis of body), prominent boss on ventral caudomesial angle. Sternum between second, third, and fourth pereiopods shallow, and setae extending from ventrolateral margins not conspicuous. First pleopod (Fig. 2a-e, g-i) as described in “Diagnosis.” In addition, proximomesial lobe of both members of pair unusually large, that of right extending mesially, caudal to and across, almost one-fourth caudal diameter of shaft of left member. < Fig. 1. Procambarus (Scapulicambarus) strenthi (all illustrations from holotype except d and i from allotype): a, Lateral view of carapace; b, Caudal view of basal podomeres of fifth pereiopod; c, Epistome; d, Annulus ventralis and adjacent steal area; e, Basal podomeres of third, fourth, and fifth pereiopods; f, Antennal scale; g, Dorsal view of carapace; h, i, Dorsal view of distal podomeres of right cheliped. 416 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 2. Procambarus (Scapulicambarus) strenthi (all illustrations from holotype except f and j from morphotype): a, b, f, g, Mesial view of left first pleopod; c, Caudal view of first pleopods; d, e, i, j, Lateral view of left first pleopod; h, Caudal view of distal part of left first pleopod. VOLUME 90, NUMBER 2 417 Uropod with both lobes of basal podomere bearing short acute spines; mesial ramus with distomedian spine situated far proximal to distal mar- gin. Allotypic female.—Differing from holotype in following respects: cer- vical tubercle distinctly spiniform; cephalic lobe of epistome subtriangular with 1 or 2 acute or subacute prominences on cephalolateral borders; fixed finger of chela (Fig. li) with 5 tubercles along opposable margin; cor- responding margin of dactyl with single row of 8 tubercles, two rows not distinguishable; single row of minute denticles on both fingers; merus of cheliped with 9 tubercles in ventromesial row, 7 in ventrolateral row, and 2 lying between distal members of rows. Annulus ventralis (Fig. ld) as described in “Diagnosis.” Sternum im- mediately cephalic to annulus with low rounded tubercles on each side of median line. Postannular sclerite broadly arched cephalically and al- most straight caudally, almost as long and four-fifths as wide as annulus. First pleopod reaching cephalic margin of annulus when abdomen flexed. (See Table 1.) Morphotypic male, form II.—Differing from holotype in following respects: acumen barely overreaching base of ultimate podomere of an- tennular peduncle; cervical tubercle not larger than others nearby; cephalic section of telson with 2 spines in caudosinistral corner and 3 in caudo- dextral; cephalic lobe of epistome subtriangular with spine near midlength of both cephalolateral margins; opposable margin of fixed finger of chela with large, more ventral tubercle situated at base of distal third of finger; opposable margin of dactyl with 7 tubercles in dorsal row and 5 in ventral; band of minute denticles on both fingers narrower; carpus of cheliped with acute tubercle on dorsomesial distal angle; merus of right cheliped with 2 spiniform premarginal tubercles dorsodistally, 13 tubercles in ventro- mesial row, 9 in ventrolateral row, and 2 lying obliquely between distal members of rows; hooks on ischia of third and fourth pereiopods much re- duced, neither reaching basioischial articulation. First pleopod (Fig. 2f, j) with juvenile basal suture marked by un- sclerotized area; shoulder on cephalic surface less well defined than in holotype and rounded distally; terminal elements all recognizable but, except for mesial process, less sharply defined; mesial process more robust and directed at right angle to shaft of appendage. Type-locality—Small stream 15 mi (24 km) W of Ciudad Valles, San Luis Potosi, Mexico. According to Dr. Strenth, there the clear, spring-fed stream, with a maximum width of about 5 m, flows through a small valley with a moderately swift current. The stream bed, with steep banks, is littered with rocks, and pools as much as 1.5 m in depth, alternate with riffles. The crayfish were obtained at night from the quieter waters just upstream from a riffle area. (This is also the type-locality of the shrimp 418 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Table 1. Measurements (mm) of Procambarus (Scapulicambarus) strenthi. Holotype Allotype Morphotype Carapace Height 11.4 9.4 9.3 Width Lele / 9.5 9.1 Total length 24.3 20.2 20.0 Postorbital length 19.1 15.9 15.6 Areola Width 0.9 0.8 1.0 Length 8.3 6.9 6.9 Rostrum Width 4.4 Sil SED Length 6.5 5.6 Sail Chela Length of mesial margin of palm 7.6 4.3 4.6 Width of palm 6.4 3:6 Sus Length of lateral margin 20.0 10.0 ball Length of dactyl 10.6 4.7 6.2 Abdomen Width 10.9 9.5 8.7 Length 26.8 ST PAllgib Palaemonetes mexicanus Strenth (1976:7), and of the gastropod Pachy- chilus apheles Thompson (1967:26).) Infesting the types was the ento- cytherid ostracod Ankylocythere toltecae Hobbs (1971:36). Disposition of types.—The holotype, allotype, and morphotype are de- posited in the National Museum of Natural History (Smithsonian Institu- tion), nos. 147725, 147726, and 147727, respectively, as are the paratypes consisting of 14 form I, 1¢ form II, and 1°. Size.—The largest specimen available is the holotype (see Table 1). Range and specimens examined.—Known only from the type-locality, 1é11, 12 with young, 14 March 1974, N.E.S., coll.; 261, 1¢1IL, 12, 29 May G74, eG ee coll Variations —Except for the presence of multiple cervical spines (3 on right and 2 on left) in the paratypic male, form I, and the single one in the paratypic female being so small as to be hardly recognizable, the variations noted in the paratypes are not noteworthy; even the numbers and dis- tribution of tubercles on the cheliped are within the limits described in the primary types. Life history notes—The allotypic female was carrying young when collected on March 14; 34 juveniles in the second instar were clinging to her abdomen when she was preserved. VOLUME 90, NUMBER 2 419 Relationships.—This crayfish is allied to the five previously described members of the subgenus Scapulicambarus: P. (S.) clarkii (Girard, 1852), P. (S.) howellae Hobbs, 1952, P. (S.) okaloosae Hobbs, 1942, P. (S.) paeninsulanus (Faxon, 1914), and P. (S.) troglodytes (LeConte, 1856). Although somewhat distantly related to them, it shares fewer characters with P. (S.) clarkii and P. (S.) troglodytes than with the other three. It may be distinguished from all of them by the deep punctations on the dorsal surface of the carapace, the broader rostrum with less convergent margins, the broader antennal scale, and especially by the features of the first pleopod. The latter is conspicuously more setose than in any of the others, and the cephalic process is unique in forming a gently curved lamelliform plate flanking the cephalic part of the central projection. In possessing a tuberculate sternum cephalic to the annulus ventralis, it re- sembles P. (S.) okaloosae and P. (S.) paeninsulanus; however, in them the tubercles overhang (ventrally) the cephalic margin of the annulus. Etymology.—This crayfish is named in honor of its original collector, Ned E. Strenth, who has donated a number of specimens of crayfishes and palaemonid shrimps to the Smithsonian Institution. Literature Cited Faxon, Walter. 1914. Notes on the crayfishes in the United States National Museum and the Museum of Comparative Zoology with descriptions of new species and subspecies to which is appended a catalogue of the known species and sub- species. Mem. Mus. Comp. Zool., Harvard Coll. 40(8):351-427, 13 plates. Girard, Charles. 1852. A revision of the North American Astaci, with observations on their habits and geographical distribution. Proc. Acad. Nat. Sci., Philad. 6:87-91. Hobbs, Horton H., Jr. 1942. The crayfishes of Florida. Univ. Fla. Publ., Biol. Scimmsemeo(2)eviy-— 179,24 pls: 1952. A new crayfish of the genus Procambarus from Georgia with a key to the species of the Clarkii Subgroup (Decapoda, Astacidae). Quart. Journ. lane Ncads Sei. o(3) 65-1745 14 figs. —. 1962. La presencia de Procambarus clarkii (Girard) en los estados de Chihuahua y Sonora, México (Decapoda, Astacidae). Anal. Inst. Biol., Mex. 383(1, 2):273-276, 9 figs. —. 1971. The Entocytherid Ostracods of Mexico and Cuba. Smithsonian Con- th ZOOl Gll—oa) ole figs. LeConte, John. 1856. Descriptions of new species of Astacus from Georgia. Proc. Acad. Nat. Sci., Philad. 7:400—402. Strenth, Ned E. 1976. A review of the systematics and zoogeography of the fresh- water species of Palaemonetes Heller of North America (Crustacea: Decapoda). Smithsonian Contrib. Zool. 228:1-27, 6 figs. Thompson, F. G. 1967. Two new species of Pachychilus from Northeastern Mex- ico. Nautilus 81(1):25-31, 2 figs. Department of Invertebrate Zoology, Smithsonian Institution, \Vashing- ton, D.C. 20560. PROC. BIOL. SOC. WASH. 90(2), pp. 420-423 PRELIMINARY ACCOUNTS OF FIVE NEW GENERA OF STOMATOPOD CRUSTACEANS Raymond B. Manning In 1968 I presented the first attempt at a revision of the stomatopod genus Squilla, then a large and heterogeneous assemblage of species. Two genera were removed from the synonymy of Squilla and eight new genera were recognized. Two of the new genera, Anchisquilla, containing four species, and Squilloides, to which six were assigned, were considered to be more heterogeneous than the remainder of the genera separated from Squilla. The study of additional material since 1968 has shown that several new genera should be recognized for species originally assigned to Anchisquilla and Squilloides. This opportunity also is taken to recognize another genus for one species previously assigned to Alima. Three of the four species originally assigned to Anchisquilla (Manning, 1968:127) are removed to new genera below, restricting Anchisquilla to the type-species, Squilla fasciata De Haan, 1844, and a second species, A. punctata Blumstein, 1974. As Moosa (1975:9) correctly pointed out, A. fasciata has fixed apices on the submedian teeth of the telson. Squilloides is restricted to two of the five species originally assigned to it (Manning, 1968:131). Since 1968 I have been able to examine the type of one species originally placed in Squilloides, Squilla minor Jurich, 1904; it proved to be a small species of Clorida, similar to but distinct from C. fallax (Bouvier, 1914). A redescription of C. minor is in preparation. Two of the other species originally assigned to Squilloides, Squilla lata Brooks, 1886 and S. gilesi Kemp, 1911, are referred to a new genus, be- low. Squilloides is restricted to the type-species, Squilla leptosquilla Brooks, 1886, and a second species originally assigned to it, Squilla tenui- spinis Wood-Mason, 1891. A planned review of families and genera of Stomatopoda from the Indo-West-Pacific region, which has been in preparation for some time, will be delayed further pending completion and publication of other studies. That review will include keys to all genera and comparative illustrations of the type-species of each genus, and the interrelationships of the higher taxa will be explored in detail. Original references for species mentioned in the text but not cited in the references below can be found in Kemp (1913). This study was supported by the Smithsonian Institution through its Research Awards and Foreign Currency Programs. I thank T. E. Bowman for his comments on the manuscript. VOLUME 90, NUMBER 2 421 Alimopsis, new genus Definition—Eye large, cornea bilobed, inner margin of eye the longer. Ocular scales separate. Carapace with median carina. Mandibular palp and 2 epipods present. Dactylus of claw with 5 teeth. Lateral process of fifth thoracic somite bilobed. Lateral processes of sixth and seventh thoracic somites not markedly bilobed. Abdomen with median carina. Submedian teeth of telson with fixed apices. Basal prolongation of uropod with inner margin crenulate. Type-species.—Squilla supplex Wood-Mason, 1875. Etymology.—The name is derived from the Greek -opsis, like, in com- bination with the generic name Alima. The gender is feminine. Remarks.—Alimopsis, which includes only the type-species, shows close affinities with Alima, as the name implies, differing mainly in the retention of the mandibular palp, the reduction of the number of epipods from 4 to 2, and the development of a median carina on the anterior 5 abdom- inal somites. The type-species is known from several localities around India (Kemp, 1913). Anchisquilloides, new genus Definition—Eye moderately large, cornea bilobed, inner margin of eye the longer. Ocular scales separate. Carapace without median carina. Mandibular palp and 4 species present. Dactylus of claw with 5-6 teeth. Lateral processes of fifth, sixth, and seventh thoracic somites single. Ab- domen with median carina. Submedian teeth of telson with movable apices. Inner margin of basal prolongation of uropod serrate. Type-species.—Squilla meneilli Stephenson, 1953. Etymology.—The name is derived from the Greek anchi-, near, in com- bination with the generic name Squilloides. The gender is masculine. Remarks.—Anchisquilloides contains only the type-species, A. meneilli (Stephenson), from Australia. The genus shows no close affinities with any other squillid genus, although the telson shape more or less resembles that found in Squilloides. Distosquilla, new genus Definition Eye large, cornea bilobed, outer margin of eye the longer. Ocular scales separate. Carapace without median carina. Mandibular palp absent, 3 epipods present. Dactylus of claw with 4 teeth. Lateral processes of fifth, sixth, and seventh thoracic somites single. Abdomen lacking me- dian carina. Submedian teeth of telson with movable apices. Basal pro- longation of uropod with row of spines on inner margin. Type-species.—Squilla miles Hess, 1865. 422, PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Etymology—The name is from the Latin disto, different, in combination with the generic name Squilla. The gender is feminine. Remarks.—Distosquilla, which includes only the type-species, shows no close affinities with any other genus of squillid. The shape of the eye is unique within the family. Lenisquilla, new genus Definition—Eye of moderate size, cornea bilobed, inner margin of eye the longer. Carapace without median carina. Mandibular palp present, usually 4 epipods present. Dactylus of claw with 6 teeth. Lateral process of fifth thoracic somite single, lateral processes of sixth and seventh somites not markedly bilobed. Abdomen without median carina. Sub- median teeth of telson with fixed apices. Basal prolongation of uropod crenulate or with row of sharp spines. Type-species.—Squilla lata Brooks, 1886. Etymology.—The name is from the Latin leni-, smooth, in combination with the generic name Squilla. The gender is feminine. Remarks.—Lenisquilla appears related to two genera with very small eyes, Clorida and Cloridopsis, resembling them in eye shape and in structure of the lateral processes of the exposed thoracic somites. It differs from both of these genera in having fixed apices on the submedian teeth of the telson. As in Clorida, the ocular scales are more or less fused into a single plate. Lenisquilla includes L. gilesi (Kemp, 1911), L. lata and L. lata spinosa (Blumstein, 1970), and L. espinosa (Blumstein, 1974). The status of the two forms described by Blumstein requires clarification; L. espinosa is almost certainly based on a juvenile, possibly a postlarva. Levisquilla, new genus Definition—Eye small, cornea bilobed, inner margin of eye the longer. Ocular scales separate. Carapace without median carina. Mandibular palp absent. Four epipods present. Dactylus of claw with six teeth. Lat- eral processes of fifth, sixth, and seventh thoracic somites single. Abdomen lacking median and submedian carinae. Submedian teeth of telson with movable apices. Basal prolongation of uropod with row of spines on in- ner margin. Type-species—Squilla inermis Manning, 1965. Etymology.—The name is from the Latin levis, smooth, in combination with the generic name Squilla. The gender is feminine. Remarks.—Levisquilla includes the type-species, known only from Japan. It resembles Clorida in general features, but differs from all species in that genus in having a smooth, flattened telson, lacking supplementary VOLUME 90, NUMBER 2 423 dorsal ornamentation. It also resembles Cloridopsis, but differs in lacking the median carina on the carapace, in having sharp marginal denticles on the telson and spines on the inner margin of the basal prolongation of the uropod. Levisquilla also differs from representatives of both of these genera in having the ocular scales distinctly separate medially. Literature Cited Blumstein, R. 1970. New stomatopod crustaceans from the Gulf of Tonkin, South China Sea. Crustaceana 18(2):218-224, figs. 1-5. —. 1974. Stomatopod crustaceans from the Gulf of Tonkin with the description of new species. Crustaceana 26(2):113-126, figs. 1-10. Bouvier, E. L. 1914. Sur la faune carcinologique de Vile Maurice. C. R. Acad. Sci. 159:698-704. Kemp, S. 1913. An account of the Crustacea Stomatopoda of the Indo-Pacific region based on the collection in the Indian Museum. Mem. Indian Mus. 4:1—217, 10 figs., pls. 1-10. Manning, R. B. 1965. Stomatopoda from the collection of His Majesty The Emperor of Japan. Crustaceana 9(3):249-262, figs. 1-2, pls. 11-12. ——. 1968. A revision of the family Squillidae (Crustacea, Stomatopoda), with the description of eight new genera. Bull. Mar. Sci. 18(1):105-142, figs. 1-10. Moosa, M. K. 1975. Notes on stomatopod Crustacea from Seribu Islands and adjacent waters with a description of a new species. Mar. Res. Indonesia LSgIWO, wires Ike Stephenson, W. 1953. Three new Stomatopoda (Crustacea) from eastern Aus- tralia. Aust. Jour. Mar. Freshw. Res. 4(1):201-218, figs. 1-4. Smithsonian Institution, Washington, D.C. 20560. PROC. BIOL. SOC. WASH. 90(2), pp. 424-439 TWO NEW SPECIES OF ELEUTHERODACTYLUS (AMPHIBIA: LEPTODACTYLIDAE) FROM NORTHEASTERN SOUTH AMERICA John D. Lynch, and Marinus S$. Hoogmoed Few Eleutherodactylus are known within the region bounded by the Rio Orino to the north and west and the Rios Amazonas and Negro to the south. Following Lynch’s (1976) species group definitions, five members of the unistrigatus group [E. grandoculis (Van Lidth de Jeude), E. inguinalis Parker, E. lacrimosus (Jimenez de la Espada), E. marmoratus (Boulenger), and E. pulvinatus Rivero], one of the auriculatus group [E. urichi (Boett- ger) ], and three of the fitzingeri group [E. gollmeri (Peters), E. gutturalis Hoogmoed, Lynch, and Lescure, and E. vilarsi (Melin)] are currently known or reported from northeastern South America. Van Lidth de Jeude’s (1904) report of E. gollmeri is based on material of one of the new species named below and that record was not addressed by Rivero (1961) in his attempt to unravel the South American records of E. gollmeri. Hoogmoed, Lynch, and Lescure (1977) and Lynch (1975) have discussed separation of the cis-andean members of the fitzingeri group described to date. In the course of fieldwork in Surinam, Hoogmoed secured series of and considerable ecologic data for the two species named below (his field- work was supported by the Netherlands Foundation for Advancement of Tropical Research (WOTRO), grants W 956-2 and W 87-78). Lynch had encountered both in museum collections and prepared preliminary de- scriptions for each. In the following accounts, specimens are identified by museum acronyms and numbers (if more than one specimen is in- cluded under a single number, the number of specimens is given paren- thetically ); the collections are as follows: American Museum of Natural History (AMNH); British Museum (Natural History), London (BM); Field Museum of Natural History, Chicago (FMNH); Museum of Natural History, The University of Kansas, Lawrence (KU); Los Angeles County Museum of Natural History (LACM); collection of Jean Lescure, Paris (LG, LGD); Museum of Comparative Zoology, Harvard University (MCZ); Rijksmuseum van Natuurlijke Historie, Leiden (RMNH); National Museum of Natural History, Washington (USNM); collection of Werner C. A. Bokermann, Sao Paulo (WCAB); and Zoologisch Museum (Instituut voor Taxonomische Zoologie), Amsterdam (ZMA). Certain abbreviations are used in the text: E-N, distance between eye and nostril; IOD; interorbital distance; and SVL, snout—vent length. Measurements were taken with dial calipers to the nearest 0.1 mm. For VOLUME 90, NUMBER 2 425 the measurements of holotypes both right and left side measurements are given if the feature is bilateral. Unless otherwise noted, all measurements refer to adult frogs; maturity in males was accepted if vocal slits were pres- ent, in females if large ova were present or if the oviducts were enlarged and convoluted. Drawings were made by Mr. W. C. G. Gertenaar and photographs by Mr. Chr. Hoorn, Jr., both of the Rijksmuseum van Natuur- lijke Historie, Leiden. Each locality is numbered (see Figs. 4, 7) and these numbers appear in parentheses in the sections listing the paratype collections. Eleutherodactylus chiastonotus, new species Holotype—RMNH 17614, an adult male, collected at Brownsberg, Brokopondo District, Surinam, 500 m, on 11 November 1968 by M. S. Hoog- moed (locality 5, fig. 4). Paratypes—(78). BRASIL, Estada Amapd: (27) Mazagao, WCAB 19177, 35968; (26) Serra do Navio, KU 140879-80, WCAB 2312-13, 3088- 89, 18166-67, 35881-82. FRENCH GUIANA: no further data, LG 1420; (21) Crique Gabrielle, LG 183; (24) Crique Grégoire, LG 15-16; (20) Eaux Clément (Montagne de Roura), LG 1230; (23) lower Matarony river, upstream from lAp- prouague, LACM 44647, 44652; (18) Monts Attachi Bacca, LG 544-45; (17) opposite Nassau mountains, RMNH 17620; (16) 10 km NE St. Laurent, RMNH 17641(2); (19) Sail, base of Mt. Galbao, LG 1408-13; (22) Saut Tortue, ORSTOM camp, LACM 44655, 44657; (25) Trois Sauts, Oyapock river, LG 1280-81, 1369. SURINAM, Brokopondo District: (5) Brownsberg, 500 m, RMNH 17613, 17615, 17616(2); Brownsberg, near Mazaronitop, 450 m, AMNH 87738-39; Brownsberg Natuurpark, MCZ 89194-97, 89211, 89214-18; (4) railway km 121, at foot of Brownsberg, RMNH 17621-22. Marowijne District: (7) Lely mountains, vicinity of Suralco camp IV, 600-690 m, RMNH 17637(3), 17639-40, 17745-46; (6) Lely mountains, 1 km S Suralco camp, V, 11 km NE airstrip, 620 m, KU 159620, RMNH 17636(2); (12) Loe Creek, 2nd camp, 120 m, RMNH 17633; (13) Loe Creek, Camp Hofwijks IV, 120 m, RMNH 17638; (15) Loée Creek, Camp Hofwijks VII, 54 km S Oelemari airstrip, 120-140 m, AMNH 90817, RMNH 17629-30, 17632, 17634, 17750; (14) Loe Creek, camp Hofwijks VIII, 56 km SSE Oele- mari airstrip, 120 m, RMNH 17631; (15) 5 km SE camp Hofwijks VII, 59 km S Oelemari airstrip, 120 m, RMNH 17635; (10) Paloemeu airstrip, RMNH 15145; (2) Patamacca, 15 km S main emplacement, RMNH 17749. Nickerie District: (9) Gonini encampment, Coeroeni river, RMNH 17618(2):; (S) King Frederick Willem IV Falls, BM 1936.9.3.15; (11) 4-5 km E Sipaliwini airstrip, RMNH 17617, 17619. Para District: (3) Afobaka road, 85 km 426 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON SSE Paramaribo, AMNH 87737. Suriname District: (1) 6 km S Blakawatra (forest service), RMNH 17642. Diagnosis.—A moderate-sized Eleutherodactylus [4 ¢ 26.7-39.7 mm SVL (x = 33.0; n = 20), 09 9°37.5=53/4 (x= 44.0) n= 14) tomithesiincen group; toes lacking webbing and lateral fringing; digits bearing pads and discs; pads of inner fingers 1.2-2.0 (x = 1.5) times digit width below pad, those of outer fingers 2.0-3.2 (* = 2.6); palmar tubercle bifid; no tarsal fold; no tubercles on heel; inner metatarsal tubercle much larger than outer; skin of dorsum finely shagreened, lacking warts; dorsolateral fold represented by row of pustules (none enlarged); tympanum prominent, its length about half eye length; choanae partially concealed by palatal shelf of maxillary arch; snout acuminate; males with vocal sac and slits; legs of moderate length, shank 51.4-64.6 percent SVL; venter cream with markings on throat; dorsum bearing prominent X-shaped mark; posterior surfaces of thigh pale tan, not patterned; limb bars as wide as interspaces, bars perpendicular to limb axis (Fig. IAB). Eleutherodactylus chiastonotus is readily distinguished from all other species of the fitzingeri group by its color pattern and markedly acuminate snout (the longest of any species in the species group). It is seemingly the northeastern South American counterpart of the Amazonian E. conspicillatus, the Venezuelan E. terraebolivaris, and the Chocoan E. achatinus, but differs from all in color pattern, snout shape, and in lacking any basal webbing of the toes. Description—Head as broad as or slightly broader than body, head longer than wide (head width 77-83% head length); head width 31.6-36.4 (x = 33.6, n = 20) % SVL in males, 32.6-36.1 (x = 34.2) 17 = 149) Simatemales: snout markedly acuminate in dorsal view, pointed and overhanging lower jaw in lateral profile (Fig. 3); snout long, E-N 91.8-123.6 (x = 110.7, n = 19) % eye length in males, 108.5-135.2 (x = 121.6, n = 14) im females; canthus rostralis sharp, straight or weakly convex; loreal region flat or weakly concave, sloping abruptly to lips; lips not flared; nostrils weakly protuberant, directed laterally; tip of snout rounded, no proboscis; inter- orbital space slightly wider than upper eyelid width, flat; upper eyelid width 67.5-117.6 (x = 90.0, n = 20) % TOD in males, 59.1-121.0 (x= 74.20; n = 14) in females; no frontoparietal fontanelle; temporal region vertical; tympanum round in males, its length 43.8-57.1 (* = 51.5, n = 20) % length; tympanum higher than long in females, its length 49.0-62.3 (x = 54.7, n = 14) % eye length; supratympanic fold prominent, thin, not obscuring upper edge of tympanum; choanae partially concealed by palatal shelf of maxillary arch when roof of mouth is viewed from directly above; choanae moderate- sized, each about as large as an odontophore; prevomerine odontophores elevated, triangular in outline (slanted or oblique in small specimens), posterior and median to choanae, separated by a distance equal to width VOLUME 90, NUMBER 2 427 Fig. 1. (A-B) Eleutherodactylus chiastonotus, holotype, RMNH 17614, 38.4 mm SVL; (C-D) Eleutherodactylus zeuctotylus, holotype, RMNH 17701, 25.4 mm SVL. of a process, bearing 3-5 teeth arranged in a transverse row across pos- terior edge of odontophore (rarely as an oblique patch); tongue longer than wide, notched posteriorly, posterior % to 75 not adherent to floor of mouth; male with large subgular vocal sac and slits. Skin of dorsum finely shagreened, becoming granular on flanks: skin of upper eyelids no more granular than that of top of head; dorsolateral fold poorly defined, granular; no supra-anal warts; skin of throat smooth, 428 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 2. Hand and foot of Eleutherodactylus chiastonotus holotype, RMNH 17614; line equals 5 mm. in males with two oblique folds near mandible; skin of venter smooth (granulations of lower flanks enroach on posterior parts of venter); discoidal folds prominent; skin on posteromedial surfaces of thigh (below vent) aerolate; forearm lacking ulnar folds or tubercles; palmar tubercle bifid . VOLUME 90, NUMBER 2 429 (Fig. 2), larger than oval thenar tubercle; supernumerary palmar tubercles prominent, smaller than subarticular tubercles; subarticular tubercles large, round, non-conical; fingers lacking lateral fringes; first finger longer than second; all fingers bearing pads, pads on inner fingers round, about 1% times width of digit below pad; pads on outer fingers about 2% times width of digit below pad; all pads bearing broad discs; fold of skin above pad not indented. No enlarged tubercles on heel; outer edge of tarsus lacking tubercles, inner edge bearing an elongate tubercle just proximal to inner metatarsal tubercle; no tarsal folds; 2 metatarsal tubercles, outer round, flat, 4 to % size of elongate (length 2 to 3 times width), non-compressed inner metatarsal tubercle; plantar surface lacking supernumary tubercles or with small tubercles at the bases of toes II-V; subarticular tubercles like those of fingers except more conical; toes lack webbing and lateral fringes; all toes bearing pads and discs, pads round, that on inner toe smaller than those on other toes; toe pads distinctly larger than pads of inner fingers, smaller than to equal in size to those of outer 2 fingers; heel of adpressed hindlimb reaches to nostril; when legs are flexed at right angles to sagittal plane, heels distinctly overlap; shank 51.4-64.6 (x = 57.1, n = 34) percent SVL. In preservative, E. chiastonotus is pale rusty-tan above with pale brown markings. The flanks are more pale than the dorsum. The canthal and supra- tympanic stripes are dark brown and fade ventrally. Labial bars are only faintly visible. The limb bars are pale brown edged with cream; the bars are about as wide as the tan interspaces and are transversely oriented. The posterior surfaces of the thighs and anal patch are medium brown. The venter is dull cream (under magnification is peppered with black); throat and upper chest bearing pale spots. Color in life——Back and legs ochre-yellow, beige or yellow-orange, with a pattern of light-brown or beige-brown bands of varying width, forming a distinct X-shaped mark on the back, chevrons or oblique bands on the back and flanks, transverse bands on the legs, an interorbital bar, 2 spots on the snout and 2 bands radiating from the eye to the upper lip. The elements of the darker pattern are bordered by a narrow black or dark- brown and a narrow pale yellow line. Some specimens (from Lée Creek and Sail) have a cream-colored vertebral stripe from the tip of the snout to the vent. Canthal and supratympanic stripe dark brown. Dorsally this stripe is bordered by a narrow pale-brown to pale-yellow line, passing over the canthus rostralis and the outer edge of the upper eyelid; ventrally there is no sharp border anterior to the tympanum; in some specimens nearly the whole loreal region and the upper lip are dark-brown. Tympanum chestnut-brown to light-brown. A dark-brown spot at the base of the forelimbs is sometimes present. Throat white, dirty-white or yellow with 430 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 3. (A-B) Eleutherodactylus chiastonotus, RMNH 17614; line equals 5 mm; (C-D) Eleutherodactylus zeuctotylus, RMNH 17701; line equals 5 mm. gray spots, belly yellowish-white. Groin, ventral, anterior and posterior surfaces of the thighs, posterior and ventral surfaces of the lower leg, and palms bright orange. The soles may be orange, gray or dark-brown. Ventral surface of forelimbs transparent flesh-colored. Iris divided in 3 parts: an upper golden part, a silvery gray, gray-brown, or gray-green lower part, both parts separated by a horizontal reddish-brown bar. Around the pupil, a narrow band, anteriorly reddish brown, posteriorly yellow. Measurements of holotype (mm).—SVL 38.4; shank 20.8-21.5; head length 15.3; head width 12.3; upper eyelid width 3.0-3.1; IOD 4.0; eye length 4.6-4.7; tympanum length 2.4-2.6; E-N 5.7-5.8; eye to tip of snout 8.0-8.3. Etymology.—Greek, in reference to the X-shaped mark (chi-like) on the dorsum. VOLUME 90, NUMBER 2 431 Fig. 4. Distribution of Eleutherodactylus chiastonotus; see list of paratypes for names of localities. Dotted lines denote 200 m contour. Distribution.—Low elevation (below 700 m) forests in northeastern Brazil, French Guiana, and Surinam (Fig. 4). Natural history—tIn Surinam, E. chiastonotus is much less abundant than is E. zeuctotylus; in Brazil and French Guiana, E. chiastonotus may be the more abundant, based on numbers of specimens available. Hoog- moed collected E. chiastonotus between 0815 and 2230 hrs. Most speci- mens were collected by day in leaf litter on the forest floor; however, the frog is a nocturnal species. Daytime collections resulted from chasing the frogs from their hiding places. Only occasional specimens were found in the daytime (early morning) sitting on leaves of low vegetation. At night, most were found on leaves and branches of herbs, shrubs, and felled trees, 3-100 cm above the ground; others were discovered on the forest floor. Eleutherodactylus chiastonotus was never found outside of the high forest. In Surinam, the frog occurs south of the coastal belt of savannas, but in some places (Patamacca and S of Blakawatra) along the southern edge of the savannas, it invades the savannas along creeks supporting gallery forests. In the Sipaliwini area, E. chiastonotus was found in the rain- 432, PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON forest to the west of the savanna but not in the forest islands surrounded by savanna. Males were observed calling as early as 1745 hrs, from elevated stations (0.5-1.0 m above ground) on thin branches of shrubs or young trees but not on tree trunks or logs. Calling males were heard and/or recorded in the months of November, January, and May. In Surinam, females containing large oviducal eggs were found in October, January, and March; females taken in July and August have only small ovarian eggs. On 28 May 1968 at 0900 hrs, several apparently recently hatched juveniles were found on leaves of Selaginella a few centimeters above the ground in high forest. Slightly larger juveniles were found on 25 June and 2 August. Thus re- production is apparently seasonal in E. chiastonotus, distributed through the short rainy season, the short dry season, and the long rainy season. The only adult female from Brazil (WCAB 35968) is gravid and was collected in December. Males collected in January had distended vocal sacs whereas those taken in October did not. The call of E. chiastonotus is a single note (Fig. 5). Eleutherodactylus zeuctotylus, new species Hylodes gollmeri: Van Lidth de Jeude, 1904:93. Holotype —RMNH 17701, an adult male collected on west slope, Vier Gebroeders Mountain, Sipaliwini, Nickerie District, Suriname, on 7 Feb- ruary 1970 by M. S. Hoogmoed (locality 34, Fig. 7). Paratypes.—(242). BRAZIL, Estado Amapa: (44) Mazagao, WCAB 19183; (43) Serra do Navio, KU 140881, WCAB 27238. Estado Amazonas: (not on map) Serra da Neblina (nr Colombia-Venezuela border), WCAB 34157. Estado Pard: (not on map) Rio Mapuera at equator, AMNH 46200; (42) Tirids (near Surinam border), WCAB 12740. FRENCH GUIANA: no further data, LG 671, 824, LDG 29; (38) Cabas- sou, near Cayenne, LG 184; near Cayenne, LG 1014-15; (37) Crique Grégoire, LG 761, 804; (39) Montagne de Kaw, LG 1492-93; (40) Saiil, LG 38, 1058-61, 1093, 1406; (41) Trois Sauts, Oyapock river, LG 1127, 1282-84, 1292. GUYANA, Rupunini District: (36) N of Acarai Mountains, W of New River, KU 69663; (not on map) Shudikar-wau, AMNH 43680. SURINAM, Brokopondo District: (11) Brownsberg, 450-500 m, AMNH 87740, RMNH 17661, 17668(2), 17673, 17682(6); Brownsberg Natuurpark, MCZ 89209-10, 89212-13; foot of Brownsberg, RMNH 17665(5); Browns- berg, near Suralco house, 500 m, RMNH 17671; Brownsberg, 2 km E Suralco house, 500 m, RMNH 17666; railway km 121, foot of Brownsberg, upper Mahami creek, RMNH 17697-98; (28) upper Gran Rio, RMNH 17689; (10) Guyana Goud Placer, RMNH 17691(7); (2) 4 km SE Kraka, VOLUME 90, NUMBER 2 433 8 - A a B 6 _-. 4 KHTZ SECONDS Fig. 5. (A) Audiospectrogram of call of Eleutherodactylus chiastonotus, holotype (RMNH 17614); tape 1968-IV A (177-202), recorded at 19 cm/sec; air temperature about 21 C. (B) Call of Eleutherodactylus zeuctotylus, holotype (RMNH 17701); tape 1970 A (120-160), recorded at 19 cm/sec; air temperature about 22 C. RMNH 17653; near Saramaca River, km 91.5, BM 1946.4.1.22. Commewijne District: (3) camp 8, Mapane area, RMNH 17688. Marowijne District: (14) Lely mountains, Suralco camp IV, 560-690 m, RMNH 17644(3), 17646( 12), 17656, 17708, 17758(4), 17759-60, 17761-64(10), 17765, 17770; (13) Lely mountains, headwaters of Djoeka creek, Suralco camp V, 610- 620 m, AMNH 90826, RMNH 17645(2), 17648(2); (35) Loe Creek, camp Hotwijks VII, 54 km S Oelemari airstrip, 120 m, AMNH 90827, RMNH 17647(2), 17650, 17654-55, 17659(2); (12) Nassau mountains, km 1.1, RMNH 17699; (30) Paloemeu airstrip, RMNH 17687, USNM_ 159027. Nickerie District: (5) Avanavero road, km 251, Paris Jacob Creek, 60 m, RMNH 17649(3), 17768(4); (7) Avanavero road, km 219, Van Ams Creek, 80 m, RMNH 17651, 17657; (4) Kabalebo River, Avanavero cataracts, 50 m, RMNH 17773; (45) Linker Kabalebo River, river camp, RMNH 17776; 15) Rechter Kabalebo River, 2nd fall, 120 m, RMNH 17757, Brokobotoval, RMNH 17751, Doublestepsval, 120 m, RMNH 17772, Grote Belgiéval, 130 m, RMNH_ 17658(4); (16) Rechter Kabalebo River, K. vallen, 180 m, RMNH 17771; 10 km S K. vallen, 180 m, RMNH 17756; (29) airstrip, Kayser mountains, FMNH 128829, 128843; SE of airstrip, Kayser Mts, RMNH 17683; (23) King Frederick Willem IV Falls, BM 1936.9.3.13-14; Lucie River, RMNH 17690(2); (25) Lucie River, 10 km NE Coeroeni airstrip, 180 m, RMNH 17766; 10-20 km N Lucie River, 200-250 m, RMNH 17767; a 434 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON (26) Eva Creek, 20 km N Lucie River, 200 m, RMNH 17754(2), 17755; (27) Vreedzaam Creek, 10 km N Lucie River, 200 m, RMNH 17753(2); (1) camp on Maratakka River, RMNH 17704; (24) encampment Tigri, New River (= upper Corantyne River), RMNH 17672; (31) upper New River (= upper Kutari Creek), BM 1936.4.2.9; (32) “New River, 230 m,” BM 1939.1.1.22-24; (6) Blanche Marie Cataracts, Nickerie River, 80 m, RMNH 17652(3), 17703; (7) Reynold’s camp (Grassalco), Mozes Creek, 90 m, RMNH 17660, 17769; 3 km S Reynold’s camp, Mozes Creek, 250 m, RMNH 17752; (33) Sipaliwini, E of airstrip, RMNH 17685, 17700, 4 km E airstrip, RMNH 17664(2), 17670, 17674, 17679-80, 17684, 17686, 17702(6); (34) Sipaliwini, Vier Gebroeders Mountain, RMNH 15168(2), 15180, 15196, 17675-77, 17681; Sipaliwini, 2 km N Vier Gebroeders Mountain, RMNH 17678(2). Saramacca District: Coppename River, RMNH 4900; (8) Coppename River, camp 1, BM 1946.4.1.3-21; (9) Coppename River, Raleigh cataracts, 50 m, AMNH 87741-43, RMNH 17662, 17663(13) 17669; (19) Linker Coppename River, ZMA 6293; Rechter Coppename River, RMNH 17695; (17) Hebiweri, RMNH 17692(2); (22) Tafelberg airstrip, 250 m, RMNH 17643(7); (21) trail from Toekoemoetoe Creek to Tafelberg, km 3, RMNH 17696; (20) Wilhelmina Mountains, transect I, RMNH 17693(4); (18) Wilhelmina Mountains, transect I], RMNH 17694. Diagnosis——A moderate-sized Eleutherodactylus [| é 6 20.4-29.6 mm SVL (x = 25.5, n = 20), 22 30:543.3 (« = 37.0, n = 32) | of the faizinzeniearomp: toes lacking basal webbing; digits bearing narrow lateral keels; digits bear- ing pads and discs, pads of inner fingers 1.2-2.0 (x = 1.5) times digit width below pad, those of outer fingers 2.2-2.9 (x = 2.5) times digit width; palmar tubercle round (not divided); no tarsal fold; no tubercles on heel; inner metatarsal tubercle much larger than outer; skin of dorsum granular; dorsolateral ridges present; tympanum prominent, round, its length about half that of eye; males with vocal sac and slits; legs of moderate length, shank 48.0-60.8% SVL; venter dusky, not patterned; dorsum mottled brown on gray ground color; posterior surfaces of thighs dark brown; limb bars transverse, slightly to distinctly wider than interspaces. No other species of the fitzingeri group has an entire (undivided) palmar tubercle. Eleutherodactylus zeuctotylus is also distinctive in having a long thumb; the long thumb and large digital pads of E. zeuctotylus are reminiscent of E. heterodactylum (Miranda-Ribeiro). The two are readily distinguished because E. heterodactylum has much narrower limb bars and smaller tympana. In habitus, coloration of the posterior thigh, and some proportions, E. zeuctotylus resembles E. fenestratus (Steindachner), E. lanthanites Lynch, and E. vilarsi (Melin). Description—Head as broad as body, slightly longer than wide; head width 36.6-39.9 (x = 38.2, n = 20) % SVL in males, 34.0-42.0 (x = 36.6, n = 32) in females; snout subacuminate in dorsal view, rounded in lateral VOLUME 90, NUMBER 2 435 profile, not extending much beyond lower jaw (Fig. 3); snout moderate in length, E-N 75.0-100.0 (x = 86.2, n = 20) % eye length in males, 70.1— 115.6 (x = 97.1, n = 31) in females; nostrils weakly protuberant, directed dorsolaterally; canthus rostralis sharp, convex; loreal region weakly con- cave, sloping abruptly to non-flared lips; interorbital space flat, no fronto- parietal fontanelle; upper eyelid width 74.2-126.1 (x = 98.8, n = 19) % IOD in males, 77.1-116.7 (x = 93.8, n = 31) in females; typanum round in males, its length 51.1-68.8 (x = 59.7, n = 20) % eye length; tympanum round to higher than long in females, its length 38.3-64.4 (x = 55.5, n = 31) % eye length; supratympanic fold not obscuring upper edge of tympanum; tympanum separated from eye by less than half tympanum length; choanae lying within palatal shelf of maxillary arch; choanae moderate-sized, each about the size of an odontophore; prevomerine odontophores tri- angular in outline, posterior and median to choanae, separated by 1% choanal widths, bearing a row of teeth across posterior edge; tongue longer than wide, weakly notched posteriorly, posterior half not adherent to floor of mouth; males with subgular vocal sac and vocal slits. Skin of dorsum finely granular to finely areolate, that of flanks more coarsely areolate, no enlarged warts; dorsolateral folds glandular; no en- larged supra-anal warts; skin of throat smooth, in males with two oblique folds near mandible; skin of venter smooth; discoidal folds prominent; skin on posteroventral surfaces of thighs (below vent) areolate; forearm lacking ulnar folds of tubercles; palmar tubercle not bifid (Fig. 6), larger than oval thenar tubercle; supernumerary palmar tubercles round, prominent; subarticular tubercles round, weakly conical; fingers bearing indistinct lat- eral keels; first finger distinctly longer than second; all fingers bearing pads and discs, pads slightly broader than long, those on inner fingers 1.2-2.0 times digit width below pad, that on finger II smallest, on outer two fingers 2.2-2.9; pads of outer fingers % to 7% tympanum length. Heel lacking enlarged tubercles; outer edge of tarsus smooth, inner edge bearing tubercle just proximal to inner metatarsal tubercle; 2 metatarsal tubercles, outer round, conical, % size of elongate (length twice width), non-compressed inner; plantar supernumerary tubercles restricted to base of toes II-IV; subarticular tubercles round, conical; toes lacking webbing; toes bearing indistinct, lateral keels; all toes bearing discs and pads, wider than long; toe pads slightly smaller than those of outer fingers; pad of toe IV largest; heel of adpressed hindlimb reaches between eye and nostril; when legs are flexed at right angles to sagittal plane, heels distinctly overlap; shank 49.5-60.8 (* = 55.2, n = 20) % SVL in males, 48.0-59.8 (+ = 54.0, n = 32) in females. In preservative, E. zeuctotylus is gray to gray-brown above with brown spotting or marbling; the body pattern varies from distinct marbling to little, if any, pattern; pattern usually diminishes on flanks; limb bars 436 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 6. Hand and foot of Eleutherodactylus zeuctotylus, holotype, RMNH 17701; line equals 5 mm. slightly wider than interspaces and transverse, but in some bars are broken up and shank is marbled; canthal and supratympanic stripes and labial bars dark brown; groin, posterior surfaces of thighs, ventral surfaces of shank dark brown; venter gray to dark brown with white flecks on throat; snout paler than dorsum, bearing pair of brown spots. In life, E. zeuctotylus is colored as follows: back orange-brown, red- brown, brown, gray brown, or greenish-gray with an indistinct darker pattern of lines and spots; flanks sometimes yellowish-green; snout, in front of eyes, paler than dorsum; upper arm of juveniles orange-brown to yellow; venter dark gray to black, throat with white spots; upper lip spotted black and white; canthal and supratympanic stripe black; tympanum chestnut- VOLUME 90, NUMBER 2 437 48° Fig. 7. Distribution of Eleutherodactylus zeuctotylus in Guiana region; see list of paratypes for names of localities. Dotted lines denote 200 m contour. brown; iris gold, divided horizontally by reddish-brown band (lower part of iris sometimes darker than upper part); groin, back of thighs, and underside of shank minium. Measurement of holotype (mm).—SVL. 25.4; shank 14.1-14.3; head length 10.8; head width 10.0; upper eyelid width 2.4-2.6; IOD 2.6; tympanum length 2.6; eye length 4.0-4.3; E-N 3.4; eye to tip of snout 5.0. Etymology.—Greek, in reference to the fused (undivided) palmar tubercle. Distribution—In Surinam, E. zeuctotylus inhabits low elevation (0- 690 m) forests. Altitudinal data are lacking for non-Surinam localities, but the altitudinal range is probably the same in Guyana, French Guiana, and northeastern Brazil (Fig. 7). In addition to the Guianas, E. zeuctotylus is known from the Serra de Neblina, in Amazonas, Brazil (near the com- mon border with Colombia and Venezuela) (WCAB collection), and from Cachoeira Santo Antonio, Estado Rondonia, Brazil (just upstream from Porto Velho) (specimens in USNM and Museu de Zoologia, Uni- versidade de Sao Paulo); these two localities are not mapped in Fig. 7. Natural history—Although this species has been collected nearly all 438 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON day and night, except during the period 0030-0645 hr, it is a nocturnal frog. Practically all specimens collected during the daytime were found in leaf-litter on the forest-floor, in heaps of fallen leaves at the base of prickly palms in leaves accumulated between buttresses or in entrances of burrows, under logs, loose pieces of bark on the ground, or under rocks. Occasionally individuals were found sitting on treetrunks, roots, or rocks in the early morning, indicating some daytime activity. Others were likely hiding among leaves or under the objects mentioned, and when startled started jump- ing around. An especially favored hiding place seems to be among leaves accumulated at the entrance of burrows; several times up to four individuals were observed to be hiding there. At night the frogs were found at quite different places, on leaves of herbs, shrubs and trees 10-300 cm above the ground, and on logs and rocks in or along creeks. The species never was found outside the forest. In most places this is high forest, but in several places (e.g., SE of Kraka), it is low forest on sandy soil or cloud-forest (e.g., in some places on Lely Mountains). In Surinam and in French Guiana the species is generally found south of the coastal savanna-belt, but there is some invasion along the southern edge where sufficient forest is present. In the Sipaliwini savanna in southern Surinam, the species occurs in forest-islands completely surrounded by savanna. The species seems to have a preference for clearings in the forest; at least males were heard calling around the edges of camp clearings throughout the country. Even recent clearings had higher concentration of calling males around them than did the surrounding forest. Males were observed calling from 1815 hr on, in the months November, December, January, February and May. They call preferably from logs or from trunks of small trees. When calling, the males may be positioned on vertical surfaces with the head pointing downward, or they may call from horizontal surfaces. Females containing large oviducal eggs were found in November, December, Jan- uary, February, March and June. Females with small ovarian eggs were found in August and September, and recently hatched juveniles in May. This means that the species is reproductively active during the short rainy season, the short dry season, and the long rainy season. From the beginning of the long dry season on, females start to form ovarian eggs which ap- parently require about 2-3 months to mature. The mating call of E. zeuctotylus is a short trill (Fig. 5). Literature Cited Hoogmoed, M. S., J. D. Lynch, and J. Lescure. 1977. A new species of Eleuthero- dactylus from Guiana (Leptodactylidae, Anura). Zool. Meded. Rijksmus. Nat. Hist. Leiden 51(3):33-41. Lynch, J. D. 1975. The identity of the frog Eleutherodactylus conspicillatus (Gimther), with descriptions of two related species from northwestern South America VOLUME 90, NUMBER 2 439 (Amphibia, Leptodactylidae). Contrib. Sci. Nat. Hist. Mus. Los Angeles Co. (272): 1—19. ———. 1976. The species groups of the South American frogs of the genus Eleuthero- dactylus (Leptodactylidae). Occ. Pap. Mus. Nat. Hist. Univ. Kansas (61): 1-24. Rivero, J. A. 1961. Salientia of Venezuela. Bull. Mus. Comp. Zool. 126:1-207. Van Lidth de Jeude, T. W. 1904. Reptiles and batrachians from Surinam. Notes Leyden Mus. 25:83-94. School of Life Sciences, The University of Nebraska-Lincoln, 68588 and Rijksmuseum van Natuurlijke Historie, Raamsteeg 2, Leiden, The Nether- lands. PROC. BIOL. SOC. WASH. 90(2), pp. 440-446 ON SOME PYCNOGONIDA OF FRENCH OCEANIA C. Allan Child There are few records of pycnogonids from anywhere in the Pacific basin other than from its surrounding continental margins. A few scattered reports appear in comprehensive coastal expedition lists (Hedgpeth, 1961; Stock, 1968), in expeditions into some of the deeper areas (Schimkewitsch, 1893; Cole, 1909), and to a few island groups such as Hawaii (Hilton, 1942) and Indonesia (Loman, 1908; Stock, 1953). This paper reports on a few specimens collected from the Tuamotu Archipelago which serve to extend the range of a species previously re- ported from there (Child, 1970), and a small series of specimens collected in New Caledonia, from which no pycnogonid records have been pub- lished. This collection of 46 specimens includes 6 species representing 4 genera. The majority are Tanystylum bredini Child, from the Tuamotus. The others are Achelia assimilis (Haswell), Achelia nana (Loman), Endeis mollis (Carpenter), Anoplodactylus batangensis (Helfer), Anoplodactylus longiformis, new species, an unidentifiable Anoplodactylus larva, and a juvenile Tanystylum species, all from New Caledonia. This collection was placed at my disposal by Dr. J. Forest of the Paris Museum Zoological Laboratory (Arthropods), and through the courtesy of Mme. Francoise Arnaud, to both of whom I express my gratitude. The specimens are deposited in the Paris Museum. Ammotheidae Dohrn, 1881 Achelia nana (Loman) Ammothea nana Loman, 1908:60-61, pl. I, figs. 1-13. Achelia nana.—Stock, 1953:300-301, fig. 14; 1954:97; 1965:14-15, figs. 1-3; 1968:16; 1974:13-14.—Utinomi, 1971:329-330. Material examined.—New Caledonia, Ile Testard Sud, Baie de St. Vin- cent, 22°00’S, 166°05’E, coll. Salvat with Mission Singer-Polignac, 11 De- cember 1961. One female. Remarks.—This specimen is so like A. nana, that I am calling it that even though the trunk is more oval than that shown in figures of the species. The palps and legs agree with these figures. This record extends the distribution of A. nana farther south in the western Pacific from its previous records in Indonesia and from Madagas- car to Japan. Achelia assimilis (Haswell) Achelia (Ignavogriphus) assimilis (Haswell)—Fry and Hedgpeth, 1969: 106 [literature, synonymy]. VOLUME 90, NUMBER 2 44] Material examined.—New Caledonia, Ile Testard Sud, Baie de St. Vin- cent, 22°00’S, 166°05’E, coll. Salvat with Mission Singer-Polignac, 18 November 1961. One female. Remarks.—This rather variable species might be expected in New Caledonia as its previously known distribution almost rings these islands. Tanystylum sp. Material examined.—New Caledonia, Ile de Pins (Kuto), algues de dragage, coll. Mme. Pruvot. One juvenile. Remarks.—This specimen looks very much like Tanystylum bredini, but only has 2 propodal heel spines. It is nearly adult. Since no Tanystylum species is known from New Caledonia, I will refrain from assigning it to any known species. Tanystylum bredini Child Tanystylum bredini Child, 1970:296-299, fig. 3. Material examined.—Tuamotu Archipelago, Iles Gambier, Teota, Banc Gaveau, 23°08’S, 134°58’W, coll. G. Seurat, from Ulva, 27 January 1904; 21 specimens. Same collecting data, 8 March 1904; 17 specimens. Remarks.—I can find no variation in the palp segmentation, lateral process tubercles, oviger, or legs. The chelifore stumps appear shorter in some specimens than in others. The species was previously known from the Society Islands and _ its distribution is extended here to the southern reaches of the Tuamotus. Endeidae Philippi, 1843 Endeis mollis (Carpenter ) Endeis mollis—Stock, 1965:31 [literature, synonymy]; 1968:59 [key]: 1975a:76; 1975b:1083-1085.—Utinomi, 1971:327. Material examined—New Caledonia, Ile de Pins (Kuto), algues de dragage, coll. Mme. Pruvot; 2 males. Remarks.—These specimens have the usual row of many cement gland pores on legs without tubercles or large spines. Phoxichilidiidae Sars, 1891 Anoplodactylus longiformis, new species aie Material examined.—New Caledonia, Ile de Pins (Kuto), 22°37’S, 167° 30’E, algues de dragage, coll. Mme. Pruvot; One male (holotype ). 442, PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON VOLUME 90, NUMBER 2 443 Description—Trunk very elongate. First 2 trunk segmentation lines complete, third line barely discernable laterally and ventrally. Lateral pro- cesses moderately long, separated by about twice their diameter, armed dorsodistally with a single small seta. Neck moderately long in relation to lateral processes. Abdomen long, erect, armed with 2 distal setae. Ocular tubercle a tall cone, rounded at apex. Eyes small, without pigment in alcohol. Proboscis thin, cylindrical, slightly bulbous toward tip. Chelifores thin; scape without tubercles, armed with 2 or 3 distal setae. Chela palm rectangular, armed with 2 or 3 small setae. Fingers strongly curved, not longer than palm, robust, overlapping when closed, armed with 3 or 4 setae on movable finger and 1 seta on fixed finger. Both fingers with 3 narrow pointed teeth. Palp buds forming slight bulges on anterior of first lateral processes. Oviger segment 3 one and one-half times length of segment 2. Terminal 2 segments together longer than fourth segment. Several setae on all but first segment and 1 short strong spine on endal surface of fifth segment. Legs moderately slender. Femur slightly longer than subequal tibiae. Femur with long dorsodistal tubercle, longer than diameter of segment. Cement glands a single row of 8 to 10 cribriform cups on very slightly raised surface. First tibia with small dorsodistal tubercle. All segments armed with a few small setae and the major segments with a single long dorsodistal seta. Propodus moderately curved, with small heel armed with 2 stout curved spines and 2 setae. Sole armed with from 5 to 7 curved spines, most with lateral setae, and a very small lamina only % the sole length. Claw slender. Small auxiliaries thin and curved. Measurements (in mm).— Trunk length (chelifore insertion to tip 4th lateral processes ) EZ Trunk width (across first lateral processes ) 0.87 Proboscis length (laterally ) 0.65 Abdomen length (laterally ) 0.32 Third leg: Coxa 1 0:22) ibia® I 1.2 Coxa 2 0.46 Tibia 2 By Coxa 3 0:31 Tarsus 0.14 Femur 1.46 Propodus 0.51 Claw 0.28 = Fig. 1. Anoplodactylus longiformis: a, Dorsal view of trunk; b, Lateral view of trunk; c, 3rd leg, with enlargement of femoral cement gland; d, Terminal segments ot 3rd leg; e, Chela; f, Oviger, with enlargement of spine. 444 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Distribution —Type-locality: New Caledonia, Southwest Pacific Ocean. Depth range: unknown. Etymology.—The proposed name reflects the discovery of another long thin form of Anoplodactylus species that belongs presumably to a group of closely related thin species. Remarks.—This species is so near Anoplodactylus longiceps that I would have assigned it to that species were it not for the 2 propodal heel spines and the 8-10 cement glands of the new species. The males of longiceps (= longicollis Williams, preoccupied) clearly have 1 heel spine and 2 cribriform cement glands raised on large swellings. This new species is related to a number of similar long tenuous forms, such as A. massiliformis, insigniformis, oculatus, and cribellatus as well as longiceps. All of these species differ from A. longiformis in their femoral cement gland count. The first two have a single gland while oculatus has 5 and cribellatus has about 15. The cement gland shape and number appear to be among the most reliable of the diagnostic characters in this genus. The new species is more closely related to A. oculatus, a species known from the British Isles, and longiceps, than to the other species mentioned. It differs from oculatus in having shorter chela fingers with teeth, in having much shorter oviger segments, in having a longer trunk and a shorter and more cylindrical proboscis. A propodal lamina is not shown for oculatus, although it may not have been recognized at the time. The new species has a very short lamina and far fewer sole spines on a curved sole. The species most closely related in both characters and geography, A. longiceps, has been found in Queensland, Australia (Williams, 1941), Western Australia (Child, 1975), and Indonesia (Stock, 1956). The dredg- ing depth is not given for the new species. Anoplodactylus batangensis (Helfer ) Anoplodactylus batangensis.—Stock, 1968:54 [literature, synonymy]; 1975a: 133; 1975b:1082-1083.—Arnaud, 1973:957.—Child, 1975:191. Material examined.—New Caledonia, Ile de Pins, (Kuto), 22°37’S, 167° 30’E, coll. Mme. Pruvot; One male. Remarks.—This circumtropical species is one of the easiest to recognize of the genus. There is apparently no other one known with a long tubular proboscis which curves dorsally. Anoplodactylus sp. Material examined.—New Caledonia, Baie de St. Vincent, Grand Tenia, 22°01’S, 165°57’E, coll. Mission Singer-Polignac, Sta. B, 9 December 1961; One larva. VOLUME 90, NUMBER 2 445 Remarks.—This specimen is without sufficient characters to assign it to any known species or even a complex of species. Literature Cited Amaud, F. 1973. Pycnogonides des récifs coralliens de Madagascar, 4. Colossendei- dae, Phoxichilidiidae et Endeidae. Téthys 4(4):953-960, 8 figs. Calman, W. T. 1923. Pycnogonida of the Indian Museum. Rec. Indian Mus. 25(3): 265-299, 17 figs. Carpenter, G. H. 1905. Pycnogonida. The marine fauna of the coast of Ireland, pt. 6. Fisheries Ireland, Sci. Investig. 1904, 4:1-8, pls. 1-3. Child, C. A. 1970. Pycnogonida of the Smithsonian-Bredin Pacific Expedition, 1957. Proc. Biol. Soc. Wash. 83(27):287—308, 5 figs. 1975. The Pycnogonida types of William A. Hilton. I. Phoxichilidiidae. Proc. Biol. Soc. Wash. 88(19):189-210, 6 figs. Cole, L. J. 1909. Pycnogonida. Reports on sci. res. exped. eastern tropical Pacific .. . “Albatross” 1904 ...1905.. ., XIX. Bull. Mus. Comp. Zool., Harvard Univ. DAMS s5=1925 pls, 1-3. Fry, W. G., and J. W. Hedgpeth. 1969. The fauna of the Ross Sea, 7. Pycnogonida, 1. Colossendeidae, Pycnogonidae, Endeidae, Ammotheidae. Mem. New Zea- land Oceanogr. Inst. 49:1-139, figs. 1-209, tables 1-16. Hedgpeth, J. W. 1961. Pycnogonida. Rep. Lund Univ. Chile Exped., 1948-49, 40. MondeWmiv. Arsskr. N: EF. Avd. 2) 57(3)21S18; 11! figs. Hilton, W. A. 1942. Pycnogonids from Hawaii. Occ. Pap. Bishop Mus. 17(3) :43- BY5), JMO) saifess Loman, J. C. C. 1908. Die Pantopoden der Siboga-Expedition. Siboga-Exped. Monogr. 40:1—88, pls. 1-15. Schimkewitsch, W. 1893. Compte-rendu sur les Pantopodes, recueillis pendant les explorations de |’Albatross en 1891. Bull. Mus. Comp. Zool., Harvard Univ. 25(2):27—43, pls. 1-2. Stock, J. H. 1953. Contribution to the knowledge of the pycnogonid fauna of the East Indian Archipelago. Temminckia 1(13):276-313, 18 figs. ——. 1954. Pycnogonida from Indo-West Pacific, Australian, and New Zealand waters. Vidensk. Medd. Dansk Naturhist. Foren. Kjobenhavn 116:1—-168, 81 figs. ——. 1956. Tropical and subtropical Pycnogonida, chiefly from South Africa. Monel, TMs 7 swiss IG rates, ——. 1965. Pycnogonida from the southwestern Indian Ocean. Beaufortia 13 (151):138-33, 46 figs. —. 1968. Pycnogonida collected by the Galathea and Anton Bruun in the Indian and Pacific Oceans. Vidensk. Medd. Dansk Naturhist. Foren. Kjobenhavn W31:7—65, 22 figs. 1974. Medio- and _ infralittoral Pycnogonida collected during the L.1.0.E. near the landbase on Nossi-Be, Madagascar. Bull. Zool. Mus. Univ. Amsterdam 4(3):11-22, 4 figs. —. 1975a. Pycnogonida found on fouling panels from the east and west coast of America. Entomol. Ber. 35:70—77, 23 figs. 1975b. Pycnogonida from the Continental Shelf, Slope, and deep sea of the Tropical Atlantic and East Pacific. Biol. Results Univ. Miami Deep-Sea Expeds. 108. Bull. Mar. Sci. 24(4):957-1092, 59 figs. 446 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON —. 1975c. Infralittoral Pycnogonida from Tanzania. Trav. Mus. d’Hist. Nat. Grigore Antipa 16:127-134, 12 figs. Utinomi, H. 1971. Records of Pycnogonida from shallow waters of Japan. Publ. Seto Mar. Biol. Lab. 18(5):317-347. Williams, G. 1941. A Revision of the genus Anoplodactylus together with a new species from Queensland. Mem. Queensland Mus. 12(1):33-39, 4 figs. Department of Invertebrate Zoology, Smithsonian Institution, Wash- ington, D.C. 20560. PROC. BIOL. SOC. WASH. 90(2), pp. 447-457 A NEW GENUS OF FLIGHTLESS IBIS (THRESKIORNITHIDAE ) AND OTHER FOSSIL BIRDS FROM CAVE DEPOSITS IN JAMAICA Storrs L. Olson, and David W. Steadman Despite the fact that parts of Jamaica are riddled with caves and sinks (see White and Dunn, 1962), knowledge of Pleistocene faunas from these sites is rather meager, particularly as compared to the remainder of the Greater Antilles. H. E. Anthony (1920a) made collections of fossils from caves in Jamaica in 1919-1920 and briefly reported on some of the mam- mals contained therein (Anthony, 1920b). Subsequent paleontological studies, based partly or entirely on Anthony’s material, include reports on lizards (Hecht, 1951), bats (Koopman and Williams, 1951), and a new endemic genus of ceboid monkey, Xenothrix (Williams and Koop- man, 1952). Up to the present there has been no published fossil record for the class Aves from Jamaica. At the American Museum of Natural History (AMNH) we recently examined some of the unsorted and unidentified specimens collected by Anthony, among which there is a fair number of birds. We have made no attempt to identify all of the avian specimens in these collections, as we understand that there is now a great deal of unstudied fossil material from Jamaica at the Florida State Museum that should be consulted by anyone attempting a more comprehensive survey of the fossil avifauna of the island. Instead, we have taken this opportunity to describe as new the single most spectacular of the fossil birds we encountered, to which description we have appended brief observations on a few of the other remains that im- pressed us as being of interest. Olson and Wetmore (1976) documented the former existence in Hawaii of a totally flightless ibis, Apteribis glenos, based on most elements of the skeleton. Although the type-material of the new Jamaican bird is less complete than that of Apteribis, these specimens can only be inter- preted as being from a second species of flightless ibis. Eight of the nine specimens were stored in four different covered boxes, each labelled “Long Mile Cave, Windsor, Trelawny, Jamaica.” Three of these specimens (two distal ends of tibiotarsi and a proximal end of a femur) were marked with Anthony's code (“J5”) for Long Mile Cave. The size of the specimens and the elements represented strongly suggest that all eight may be from a single individual. The shaft of a humerus, apparently referable to the same species, we discovered in a box labelled “Fowl House Cave,” which is also in Windsor, Trelawny. The coloration and preservation of this specimen is so nearly like that of the others (a resemblance heightened by the fact that all nine specimens had at some 448 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON time been treated with shellac) that we strongly suspect that this humerus was actually collected in Long Mile Cave, and was erroneously introduced into the container of material from Fowl House Cave during the more than fifty years since its collection. Anthony’s Jamaican fossil sites have been discussed in general by Koop- man and Williams (1951) and Long Mile Cave in particular by Williams and Koopman (1952), this being the type-locality of Xenothrix mcgregori. Anthony's field notes (Williams and Koopman, 1952) indicate that the deposits in Long Mile Cave consisted of a small Amerindian midden over- lying an accumulation of “yellow limestone detritus” that evidently ante- dated human occupation. Anthony recorded that the primate remains came from this lower layer but unfortunately no such note was made of the ibis bones; therefore, all one can say of their age at this point is that they are late Quaternary. Each of the specimens of the new ibis was first compared with a synoptic series of skeletal elements containing all modern non-passerine families of birds. The high, pointed intercotylar knob of the tarsometatarsus and the broad, perforate procoracoid process separate the new Jamaican species from most of the families and orders to be expected in the Antilles such as Pelecaniformes, Anseriformes, Falconiformes, Galliformes, Columbiformes and the various orders of small land birds. In the Gruidae and Aramidae (Gruiformes) the shape of the hypotarsus is quite different from that of the fossil and the hypotarsus contains closed canals, which the fossil does not. The same is true for the Ardeidae. In the Phoenicopteridae the inter- cotylar knob is much wider than in the fossil; in the Ciconiidae it is much narrower and constricted at the base, in addition to which the procoracoid is not perforate. There is no group to which the Jamaican fossils show a greater similarity than the Threskiornithidae. Material of the following species of Threskiornithidae was used in the comparisons below: complete skeletons of Threskiornis aethiopicus, T. melanocephalus, T. (“Carphibis”) spinicollis, Nipponia nippon, Hagedashia hagedash, Harpiprion caerulescens, Theristicus caudatus, T. melanopis, Mesembrinibis cayennensis, Eudocimus albus, E. ruber, Plegadis falcinel- lus, P. chihi, Platalea leucorodia, and P. (“Ajaia”’) ajaja; partial skeletons of Theristicus branickii, Platalea minor, and P. regia; tibiotarsi and tarsometa- tarsi removed from skins of Cercibis oxycerca and Phimosus infuscatus; all the appropriate fossil elements of Apteribis glenos. Descriptive ter- minology follows that of Howard (1929). Threskiornithidae Xenicibis, new genus Type.—Xenicibis xympithecus, new species. Diagnosis.—A large, heavy-legged ibis with wing and pectoral girdle VOLUME 90, NUMBER 2 449 proportionately quite small. Differs from the other genera of Threski- ornithidae examined as follows: tarsometatarsus with hypotarsus simple, consisting of 2 calcaneal ridges separated by a deep groove, the internal ridge being much shorter than the external one; tibiotarsus with very wide anterior intercondylar fossa, reduced supratendinal bridge, very large outer cnemial crest extending far laterally, inner cnemial crest displaced much farther laterally than in other ibises; femur with head larger and more proximally oriented, antero-proximal portion of shaft greatly exca- vated and flattened; phalanx | of digit IV very stout; coracoid with sternal end thin and weak, area of sterno-coracoidal impression nearly convex and lacking thin diagonal ridges, internal distal angle slender and exca- vated on ventral surface. Etymology.—Gr. xenikos, strange, and ibis, f. a wading bird, an ibis. Xenicibis xympithecus, new species Figs. 1-3 Holotype.—Proximal half of left tarsometatarsus, vertebrate paleon- tological collections AMNH 11006. Collected 16 or 17 January 1920 by H. E. Anthony at Long Mile Cave, Windsor, Trelawny Parish, Jamaica. Late Quaternary. Measurements of holotype.—Proximal width 18.6 mm, proximal depth through hypotarsus 17.5, width and depth of shaft 30 mm from apex of intercotylar knob 9.6, 6.4. Paratypes.—Eight other specimens, all of which, with the possible excep- tion of the humerus noted above, have the same data as the holotype and are in the AMNH vertebrate paleontological collections. Nearly complete right coracoid, AMNH 11008; left coracoid lacking sternal end, AMNH 11013; shaft of right humerus, AMNH 11031; proximal end of left femur, AMNH 11005; imperfect proximal end of left tibiotarsus, AMNH 11009: distal *% of left tibiotarsus, AMNH 11007; distal *s of right tibiotarsus, AMNH 11004; pedal phalanx 1 of left digit IV, AMNH 11011. Measurements of paratypes——Coracoid AMNH 11008—greatest length 50.0 mm, length from internal distal angle 47.8, least width of shaft 7.9, least depth of shaft 3.3; humerus—width and depth of shaft at distal end of attachment of latissimus dorsi posterioris 6.2, 6.4; femur—proximal width 22.6, transverse diameter of head 10.1, proximal depth 17.5, width and depth of shaft 40 mm from proximal end 8.0, 8.7; tibiotarsus AMNH 11004—distal width 15.6, distal depth 15.6, least width of shaft 8.4, least depth of shaft 5.6, length of internal condyle 9.4; tibiotarsus AMNH 11009—proximal width through outer cnemial crest 28.2; phalanx 1 of digit IV—length 21.7, proximal width 7.6, least width 4.2, distal width 5.7. Diagnosis.—As for the genus. 450 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 1. Xenicibis xympithecus, new genus and species. a, Holotype proximal end of left tarsometatarsas AMNH 11006, anterior view; b, Same, posterior view; c, Same, medial view; d, Same, proximal view; e, Proximal end of left femur AMNH 11005, anterior view; f, Same, posterior view; g, Pedal phalanx 1 of left digit IV AMNH 11011, dorsal view; h, Same, ventral view. All figures natural size except d = 1.5x. VOLUME 90, NUMBER 2 A451 Fig. 2. Xenicibis xympithecus, new genus and species. a, Distal end of right tibiotarsus AMNH 11004, anterior view; b, Same, distal view; c, Distal end of left tibiotarsus AMNH 11007, anterior view; d, Proximal end of left tibiotarsus AMNH 11009, posterior view; e, Same, anterior view; f, Same, proximal view. All figures natural size except b and f = 1.5. Arrows indicate the laterally displaced position of the internal cnemial crest. Etymology.—Gr. xyn, a variant of kyn or the more usual syn (from which L. cum), meaning “with,” “along with,” or “in company with,” and Gr. pithekos, monkey. The specific name is a masculine noun in apposi tion and is an allusion to the possibility that the flightless ibis and Xeno 452 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 3. Xenicibis xympithecus, new genus and species. a, Right coracoid AMNH 11008, ventral view; b, Same, dorsal view; c, Shaft of right humerus AMNH 11031, palmar view; d, Same, anconal view. All figures natural size. thrix mcgregori existed as contemporaries in the vicinity of Long Mile Cave, where both were interred. Description.—Tarsometatarsus with intercotylar knob much higher than in Theristicus, Harpiprion, Mesembrinibis, or Phimosus, and more ver- tically oriented than in Cercibis; internal cotyla wider and projecting far- ther medially than in Platalea; prominences at posterior margins of both cotylar surfaces better developed than in other ibises, in some of which these may be lacking; hypotarsus simple, with 2 calcaneal ridges and a deep groove, differing from all other genera examined in having the internal ridge much shorter than the external ridge; external ridge simple and narrow, differing from Nipponia, Harpiprion, Theristicus, and Cercibis, in which the external portion of the hypotarsus is wider and bears 2 ridges; antero-lateral surface just distal to external cotyla more deeply excavated than in other ibises examined, although Eudocimus and Threskiornis ap- proach this condition; proximal portion of postero-medial surface of shaft much less excavated than in Hagedashia, Harpiprion, Mesembrinibis, or Phimosus. Tibiotarsus with outer cnemial crest larger, more thickened, and more laterally protruding than in any other genus of ibises, being approached only by Apteribis; inner cnemial crest displaced much farther laterally than in other ibises and more similar to the condition in kiwis (Apteryx); supratendinal bridge more reduced than in other genera examined, being composed of two triangular projections of bone whose apexes barely VOLUME 90, NUMBER 2 453 coincide over the midline of the tendinal canal in AMNH _ 11007, while in AMNH 11004 the medial side of the canal shows no sign of an excres- cence or attachment of bone, the bridge therefore having been incomplete: anterior intercondylar fossa much wider than in other Threskiornithidae and internal ligamental prominence more reduced than in other genera except Hagedashia and Apteribis; resembles typical ibises and differs from spoonbills (Platalea) in having distal third of shaft broader and shallower and the distal articulation in distal view deeper and narrower. Femur with head more protrusive above level of iliac facet, with pos- terior margin of said facet being more produced over the shaft than in other genera examined; antero-proximal surface of shaft much more deeply excavated than in other ibises, resulting in a very prominent trochanteric ridge; attachment of round ligament much deeper and situated more an- teriorly than in other genera examined, being accented by a postero-lateral prominence. Phalanx 1 of digit IV much stouter than in other ibises, with its length being 2.8 times the proximal width, as opposed, for example, to 3.6 times the width in Nipponia. Coracoid with sterno-coracoidal “impression” nearly convex and without raised striae, rather than deeply concave and with striae, as in all other ibises; sternal end very thin, internal distal angle slender and excavated on ventral surface, unlike other ibises; intermuscular ridge on ventral sur- face of shaft less distinct than in other forms; area beneath furcular facet not extensively pneumatized, as in the other taxa examined, except Apter- ibis; furcular facet sloping ventro-sternally, as in Apteribis, this facet being perpendicular to the axis of the shaft in other genera. Humerus proportionately small, with deltoid crest thicker and more palmarly rotated than in any ibis except Apteribis; shaft at distal end of attachment of M. latissimus dorsi posterioris deeper than wide, being ap- proached only by Apteribis. Remarks.—That Xenicibis xympithecus was flightless can scarcely be doubted. The elements of the hindlimb indicate a bird larger and heavier than any known ibis except Thaumatibis gigantea, while the coracoid is slightly smaller than, for example, in Threskiornis spinicollis. Furthermore, the peculiar conformation of the coracoid is indicative of reduced muscular development, and the structure of what remains of the deltoid crest on the one available fragment of humerus is that typical of a variety of flightless birds. The Hawaiian flightless ibis, Apteribis glenos, differs markedly from Xenicibis and must have evolved independently. Most of the known elements of Xenicibis differ so much trom other genera that the relationships of the Jamaican bird are not at all clear. Only the proximal end of the tarsometatarsus seems to afford some clues, being most similar to Eudocimus and quite distinct from such endemic South American genera as Mesembrinibis, Phimosus, and Harpiprion. 454 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON The tarsometatarsus of Xenicibis agrees with that of Eudocimus in the shape of the intercotylar knob, in the deep excavation on the antero- lateral surface distal to the external cotyla (greater than in other ibises, including Plegadis) and in the greater development of the prominence posterior to the external cotyla. Xenicibis differs from Eudocimus in the construction of the hypotarsus, in its somewhat deeper anterior groove with more widely spaced proximal foramina, and in the greater excavation of the postero-medial surface of the proximal end of the shaft. In proximal view Xenicibis has a distinct anterior projection of the external cotyla and a posterior projection of the internal cotyla, both absent in Eudocimus, and the neck of the hypotarsus is less constricted. In the other known elements of the skeleton, Xenicibis differs from Eudocimus in the char- acters mentioned in the generic diagnosis. Eudocimus and Plegadis are the only genera of true ibises found in North America and the Antilles. Although Eudocimus might possibly have given rise to Xenicibis, the nature of the present material of the latter does not afford any reasonable assurance of this. Accipitridae Genus and species indeterminate Also from Long Mile Cave is an ungual phalanx of a large raptor (AMNH 11010). Although the specimen lacks the ventral articular knob (Fig. 4a), enough remains of the articulation to show that this claw is not that of an owl (we compared it specifically with claws of Tyto ostologa, a giant extinct barn owl from Hispaniola). Its flattened ventral surface shows that it is not the claw of an osprey, Pandion haliaetus. The speci- men is too large (dorsal chord 32.6 mm) for Buteo jamaicensis, the largest accipitrid known from Jamaica. It is not as heavy as those in most of the New World hawks and eagles with which we compared it, but in size and slenderness it does seem rather similar to the claws of Spizastur melanoleucus, a species that now ranges from southern Mexico to Argen- tina. Even if complete, we would not consider this specimen as being sufficiently diagnostic to allow identification to genus. However, it is an indication of a large raptor now vanished from Jamaica. Large extinct or extirpated species of accipitrids have previously been recorded in the West Indies from the Bahamas (Wetmore, 1937; Brodkorb, 1959) and Cuba (Wetmore, 1928; Arredondo, 1976). Strigidae Speotyto ct. cunicularia Among the material from Dairy Cave, Dry Harbour, St. Ann Parish (see Koopman and Williams [1951] and Hecht [1951] for information on this VOLUME 90, NUMBER 2 45; Ol Fig. 4. a, Accipitridae, genus and species indeterminate, ungual phalanx AMNH 11010; b, left humeri of Siphonorhis americana (left) and S. brewsteri (right) to show difference in size. All figures natural size. site) we noted two specimens of a strigid owl much smaller than either Pseudoscops grammicus or Tyto alba, the only two owls now known from Jamaica. The proximal end of a tarsometatarsus (AMNH _ 11022) and the distal end of a tibiotarsus (AMNH 11033) agree in size and details with the Burrowing Owl, Speotyto cunicularia, and differ considerably from Otus and greatly from Glaucidium, the only other genera of small owls now occurring in the West Indies. Speotyto is known historically to have been resident in the Antilles only in the Bahamas, Hispaniola, Antigua, Nevis, St. Christopher, and Marie Galante, all the Lesser Antillean popula- tions having been exterminated within the last century. The former existence of Speotyto in Jamaica assumes considerable significance because unpublished paleontological studies by Olson and others indicate that these owls were once present on several other Antillean islands where they no longer are found. Bones of Speotyto will probably prove sufficiently abundant in Jamaican deposits to allow a more definitive assessment of the specific or subspecific characters of the Jamaican bird than the frag- ments we have seen will permit. For this reason we only tentatively refer it to S. cunicularia. Nyctibiidae Nyctibius griseus The Common Potoo, Nyctibius griseus, occurs from southern Mexico to Argentina and in the West Indies on Jamaica and Hispaniola only. The sole fossil record for the entire family is the occurrence of a Nyctibius in a Pleistocene cave deposit in Brazil listed by Brodkorb (1971) under N. griseus but only tentatively identified as this species by the original author 456 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON (Winge, 1887). It seems worthwhile, therefore, to note a left coracoid of Nyctibius griseus (AMNH 11012) among the specimens from Long Mile Cave, the second fossil record for the family and the first for the West Indies. Caprimulgidae Siphonorhis americana This distinctive nightjar, endemic to Jamaica, is now extinct. Greenway (1958) states that only three skins of this species are known, but he evi- dently did not include the one in the National Museum of Natural History (USNM 22109, collected ‘near Spanishtown’ by W. T. March). The only other species of Siphonorhis is the smaller S. brewsteri of Hispaniola. Peters (1940), who perhaps never saw a specimen of the Jamaican bird, listed S. brewsteri as a subspecies of S. americana. In this judgment he has rightfully been ignored by most subsequent authors, as the two forms differ in coloration and are very different in size. In the fossil material from Dairy Cave, we found three caprimulgid humeri that most probably pertain to S. americana, as they are very dif- ferent from Chordeiles minor, the only other caprimulgid on Jamaica. The most complete of these humeri (AMNH 11020) is 31.5 mm long as com- pared to 25 mm in a specimen of S. brewsteri (USNM uncatalogued) from a cave in Haiti (Fig. 4b). Acknowledgments For the kind assistance that led to our discovery of Xenicibis we are much indebted to Karl F. Koopman of the Department of Mammalogy, American Museum of Natural History. To Malcolm C. McKenna and other members of the staff of the Dept. of Vertebrate Paleontology at AMNH we are grateful for the loan of specimens and many other courtesies. For reading and commenting on the manuscript we thank John Farrand, Jr., Karl F. Koopman and Larry D. Martin. Arnold D. Lewis carefully re- moved layers of ancient shellac from the specimens of Xenicibis to make them more suitable for photography. The photographs were taken by Victor E. Krantz. Literature Cited Anthony, H. E. 1920a. . ee yi TUES RO Ce 2 How ae t 1 J = eg =e = i / } Vf es eat Hee. ‘. ee We 1 an Heh t N 7 4. ee MA ag hae och A - : ie i x ' 7 ee =; SU : a J { i ; a 7 - v ? - ni - I ; a y ‘, ~~ D ' : ca 2 = : i : Bai : t ‘I : at ; ix . \ f \ fas a h a ; ; J if = J : = : i : uy cs ‘ f if ~ : . oe ae el i ; ; in a 5 > — =i ‘ i =a : \ 4 P : i > = = ; ; 1 = = , age || rs P i = re i \ — i : : 7 @ ' é o Ys « = 1 oe . r : ; ; a » \ me a mi . a 1 u re) ? ' : i lcci : 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, Washing- ton, D.C. 20560. Review.—One of the Society’s aims is to give its members an opportunity for prompt publication of their shorter contributions. 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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 Redescription of Salmincola longimanus Gundrizer, 1974 (Copepoda: Lernaeopodi- dae). Z. Kabata A new species of frog of the genus Eleutherodactylus (Amphibia: Leptodactylidae) from the Cockpit Country of Jamaica. Ronald I. Crombie The synonymy and distribution of the estuarine Hypaniola florida (Hartman) from the east coast of the United States (Polychaeta: Ampharetidae). Marian H. Pettibone Scale organs on the head of Leptotyphlops (Reptilia, Serpentes): a variational study. Braulio Orejas-Miranda, George R. Zug, Daniel Y. E. Garcia, and Federico Achaval Distributional and nomenclatural notes on some Gelastocoridae in the collection of the Bernice P. Bishop Museum (Hemiptera). E. L. Todd Vargula tsujii, a new species of luminescent Ostracoda from Lower and southern California (Myodocopa: Cypridininae). Louis S. Kornicker, and James H. Baker A new species of Aphodius (Coleoptera: Scarabaeidae) from sand dunes in Chihuahua, Mexico. Robert D. Gordon Studies of tropical American Leguminosae—VIII. Richard S. Cowan The geckoes (Sauria, Gekkonidae) of the genus Sphaerodactylus of the Dominican Peninsula de Barahona, Hispaniola. Albert Schwartz Eudrilin earthworms, including a new genus, from eastern Nigeria. A. O. Segun Bonaducecytheridae, a new family of cytheracean Ostracoda, and its phylogenetic significance. K. G. McKenzie Ophiomastix koehleri, a new ophiocomid brittlestar (Echinodermata: Ophiuroidea) from the western Indian Ocean. Dennis M. Devaney Notes on the biology of the pontoniine shrimp Lipkebe holthuisi Chace, with a description of the male. J. Kevin Shaw, Richard W. Heard, Jr., and Thomas S. Hopkins The identity of Lewis’ marmot, Arctomys lewisii. Robert S. Hoffmann The milliped genus Croatania (Polydesmida: Xystodesmidae). Rowland M. Shelley Hyphessobrycon socolofi, a new species of characoid fish (Teleostei: Characidae) from the Rio Negro of Brazil. Stanley H. Weitzman A new species of characoid fish, Hyphessobrycon diancistrus, from the Rio Vichada, Orinoco River drainage, Colombia, South America (Teleostei: Characidae). Stanley H. Weitzman Conchoecia convexa, a new species of halocyprid ostracod from the Caribbean Sea and Gulf of Mexico. Georgiana B. Deevey A new crawfish of the genus Hobbseus from northeast Mississippi, with notes on the origin of the genus (Decapoda, Cambaridae). J. F. Fitzpatrick, Jr. Studies of Neotropical caddisflies, XXI. The genus Lepidostoma (Trichoptera: Lepidostomatidae). - Oliver S. Flint, Jr., and Joaquin Bueno-Soria Marine chloromonads: more widely distributed in neritic environments than pre- previously thought. Alfred R. Loeblich III, and Karen E. Fine A redescription of Oithona dissimilis Lindberg 1940 with a comparison to Oithona hebes Giesbrecht 1891 (Crustacea: Copepoda: Cyclopoida). Frank D. Ferrari A new crayfish (Decapoda: Cambaridae) from San Luis Potosi, Mexico. Horton H. Hobbs, Jr. Preliminary accounts of five new genera of stomatopod crustaceans. Raymond B. Manning Two new species of Eleutherodactylus (Amphibia: Leptodactylidae) from north- eastern South America. John D. Lynch, and Marinus S. Hoogmoed On some pycnogonida of French Oceania. C. Allan Child A new genus of flightless ibis (Threskiomithidae) and other fossil birds from cave deposits in Jamaica. Storrs L. Olson, and David W. Steadman 189 194 205 209 214 218 232 237 243 255 263 274 284 291 302 326 348 358 367 375 388 Proceedings of the BIOLOGICAL SOCIETY of WASHINGTON Volume 90 17 October 1977 Number 3 THE BIOLOGICAL SOCIETY OF WASHINGTON 1977-1978 Officers President: Richard S. Cowan Secretary: W. Duane Hope Vice President: Clyde F. E. Roper Treasurer: Oliver S. Flint, Jr. Custodian of Publications: John H. Miles Elected Council J. Laurens Barnard Raymond B. Manning Ronald I. Crombie John H. Miles Frank D. Ferrari PROCEEDINGS Editor: Austin B. Williams Coeditor: C. W. Hart, Jr. Associate Editors Classical Languages: George C. Steyskal Invertebrates: Thomas E. Bowman Plants: David B. Lellinger Vertebrates: Leslie W. Knapp 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 Institution, Washington, D.C. 20560. The Proceedings of the Biological Society of Washington is issued quarterly. Manuscripts, corrected proofs, editorial questions should be sent to the Editor, Biologi- cal Society of Washington, National Museum of Natural History, Smithsonian Institution, Washington, 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 Application to mail at second class postage rates is pending at Washington, D.C., and additional mailing office. PROC. BIOL. SOC. WASH. 90(3), pp. 459-482 THE SORICIDAE OF TAIWAN E. W. Jameson, Jr., and Gwilym S. Jones The Soricidae of Taiwan are represented by Suncus and Crocidura in the Crocidurinae and Anourosorex, Chimarrogale, and Episoriculus in the Soricinae. These shrews were collected at various localities on Taiwan and we have made some observations concerning their habits, distribu- tion and systematics. Little has been published about Taiwan soricids other than checklists (e.g. Kuroda, 1938; Chen, 1956) and fragmentary records. Only Tanaka (1936) considered the distribution and habitats of the insectivores. Kuroda (1952) discussed the zoogeographical affinities of the soricids as well as the other species on the island. Material was collected during visits from 1963 to 1969 in Taipei Hsien, Taichung Hsien, Nantou Hsien, Hualien Hsien, and Chiai Hsien. Further information about the species was obtained from the collection in the U.S. National Museum of Natural History (USNM), Washington, D.C. and from the literature. Unless otherwise indicated, distributional records and measurements are based on our collections (deposited in the USNM and the University of California, Davis), those of the late Professor Sherman A. Hoslett (deposited in the University of California, Davis), and those in the USNM. We follow Repenning (1967) in the allocation of genera to sub- families. All measurements, taken with dial calipers, are in millimeters (mm). Most are self explanatory. External measurements are those from the speci- men tags; hind foot lengths include the claw. Palate length is from the front edge of the premaxilla, viewed ventrally, to the posterior edge of the palatine. Length of the upper tooth row (UTR) is from the front edge of the incisor to the posterior edge of M3. Height of the braincase is from the highest point of the sagittal crest to the basisphenoid-basioccipital suture. Length of the lower tooth row (LTR) is from the posterior edge of of m3 to the anterior edge of pml. Height of rostrum is from the alveolus of I3 to the top edge of the rostrum. Length of mandible is from the tip of the incisor to the tip of the articular condyle. PM1 and I] is the greatest length of the respective cingulum, viewed laterally. Terminology follows Meester (1963). The colors were measured under a common light source with a Munsell Soil Color Chart; when cited, the Munsell terms are capitalized. Comparative specimens were borrowed from the American Museum of Natural History, New York (AMNH), USNM, National Science Museum, 460 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Tokyo (NSMT), Field Museum of Natural History, Chicago (FMNH), British Museum (Natural History), London (BMNH), Rijksmuseum van Natuurlijke Historie, Leiden (RMNH), and B. P. Bishop Museum, Honolulu (BPBM). Crocidura attenuata tanakae Kuroda Sorex sp. Swinhoe, 1864:382; Aoki, 1913:270, 1914:30. Crocidura attenuata Milne-Edwards, 1872 Chen, 1948:43; Jones, Lim and Cross, 1971:270; Jones, 1975:185. Chodsigoa sodalis Thomas, 1913 Kuroda, 1935:287 (misident.). Crocidura sp. Tanaka, 1936:312; Harrison and Audy, 1951:179. Crocidura tanakae Kuroda, 1938 Kuroda, 1938:81, 1940:178; Okada, 1938:2; Chen, 1956:48; 1969:349. Crocidura attenuata tanakae Kuroda, 1938 Kuroda, 1952:286; Ellerman and Morrison-Scott, 1951:83, 1966:83; Jones, Huang and Chang, 1969:49; Kuntz and Dien, 1970:33. This shrew was first described from Taiwan as a new species, C. tanakae Kuroda, 1938. The subspecies is found from sea level to at least 1,225 m at Wushe, Nantou Hsien. It occurs commonly in cane fields west of the city of Taichung in loose, sparse litter under shrubs and small trees but ap- pears to be more common in uncultivated rocky patches amid dense tangles of bushes and bamboo. C. attenuata was the most commonly caught mammal in an overgrown cultivated field near Wushe. This shrew was also collected in a cornfield at Wushe, in a weed field near Wanta, in secondary streamside growth near Meichi, and in seaside scrub vegetation and cultivated fields near Tamsui. Females with two and three embryos were found in February. The left upper unicuspids of USNM 358668 from Tamsui and the third upper unicuspid of USNM 358662 from Wushe are all missing with no apparent alveoli. Measurements.—Total length males (6 specimens) mean = 136.5 mm (range = 129-142), females (7) 133.4 (126-141); tail males (7) 52.9 (49-57), females (8) 51.8 (50-55); hind foot males (7) 13.3 (12-14), females ) 13.7 (13-15); greatest length of skull males (6) 20.5 (19.6-21.1), females 20.5 (20.2-21.0); palatal length males (6) 8.6 (8.0-8.9), females (8) (8.4-8.7); greatest breadth braincase males (6) 9.2 (8.8-9.4), females (8) 9.1 (8.9-9.4); length upper tooth row males (6) 9.2 (8.8-9.5), females (9) 9.0 (8.9-9.2); greatest maxillary width males (7) 6.6 (6.3-6.9), females (9) 6.6 (6.4-6.8). (9 ( 8.) y VOLUME 90, NUMBER 3 461 Specimens examined (17 total).—Taipei Hsien: Tamsui 25°10’N, 121°26’E (USNM 358668). Taichung Hsien: Ma-an Liao, 6 mi S Tung Hsi 24°15’N, 120°49’E (USNM 294129). Nantou Hsien: 3 km E Wushe 24°02’N, 121°08’E (USNM 358109, 358110, 358660, 358111); 1.7 km W Wushe (USNM 308661); Taita Agricultural Farm, 5.6 km E and 1.4 km N Wushe (USNM 308662); Lung Yuen Bridge, Wushe (USNM 332814); 2 km E Wanta 23°58’N, 121°08’E (USNM 358112); Meichi, 4.5 km SW Wushe 24°0I’/N, 121°06’E (USNM 358113, 358665, 358667); Meichi, 3.56 km SW Wushe (USNM 358663, 358664); Meichi, 2.5 km SW Wushe (USNM 358114, 308666). Literature records.—Taipei Hsien: Taihoku (now Taipei) 25°03/N, 121°30’E (Tanaka, 1938). Taichung Hsien: Taichu-shi (now Taichung) 24°09’N, 120°41’E (Kuroda, 1938). Chiayi Hsien: Kagi (now Chiayi) 23°29’N, 120°27’E (Kuroda, 1938; Tanaka, 1938). Nantow Hsien: Horigai (now Puli) 23°58’N, 120°57’E (Kuroda, 1952); Suiriko (now Shui-Li- Keng) 23°49’N, 120°51’E (Kuroda, 1938); Shoho-sha (now Hsiao-Pu) 24°01’N, 120°57’E (Kuroda, 1938). Crocidura horsfieldi kurodai, new subspecies Crocidura horsfieldi tadae Tokuda and Kano, 1936 Kuntz and Dien, 1970:33. Crocidura sp. Jones, Lim and Cross, 1971:271. Holotype.—Adult female, skin and skull, USNM 358115, from Linkou, Taipei Hsien, Taiwan; obtained January 1969 by Gwilym S. Jones, original no. T1451. First upper incisors broken. Diagnosis.—Distal ends of dorsal hairs vary from Dark Reddish Brown to Very Dark Brown with Dark Gray bases. Differences in color from base to tip cause grizzled appearance. Ventral fur ranges from Gray to Very Dark Gray and is same color at base as at tip. Dorsum and venter evenly colored throughout. Rather sharp demarcation between dorsal and ventral colors. Tail unicolored in type, USNM 358115, and in immature USNM 311473. USNM 332813, however, has slightly bicolored tail. All have typical bristles on basal % of tail. Hind feet gray brown, although USNM 332813 has more gray and less brown than others. Vibrissae of all three specimens are brown, Dark Brown on 358115, Medium Brown on other two specimens, and evenly colored throughout. Length of palate short, rostrum relatively massive; upper tooth row relatively short; brain- case relatively narrow; posterior edge of palatine narrow. Measurements.—Holotype plus one unsexed skin. Total length mean 104 mm (range = 97-110); tail 44 (42-46); hind foot 12 (12); palatal length 6.3 (6.1-6.5); greatest breadth braincase 7.7 (7.7); length upper tooth 462 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON row 7.1 (7.0-7.2); greatest maxillary width 5.2 (5.1-5.3); least interorbital breadth 3.7 (3.7); height of braincase 4.2 (4.2); length of lower tooth row 5.2 (5.0-5.3), length of PM4 at cingulum 1.8 (1.7-1.8), width of posterior edge of palatine 2.1 (2.1), length of nasals 4.8 (4.8); height of rostrum at I3 1.8 (1.7-1.9); length of I1 at cingulum 1.2 (1.0-1.3); distance from infra- orbital foramen to anterior point of premaxilla 4.4 (4.4). Comparison.—Crocidura horsfieldi kurodai is a medium sized C. hors- fieldi, the external measurements being closest to C. h. indochinensis from Burma (Table 1). It is smaller than C. h. watasei, Ryukyu Islands, and larger than C. h. tadae, Lan Yu Island, Taiwan and C. h. ssp., Fukien. There is general gradation in size from the smallest, C. h. tadae, through C. h. kurodai, C. h. indochinensis, C. h. horsfieldi to C. h. watasei. The type of C. h. kurodai and USNM 311473 are closest in dorsal color to C. h. watasei, whereas USNM 332813 is closer to C. h. horsfieldi and those C. h. indochinensis from Burma. C. h. tadae (NSMT 4395) has a Dark Reddish Brown dorsum, but unlike C. h. kurodai and the other sub- species studied, which have a grizzled color due to bicolored dorsal hairs, it has a solid rusty color due to unicolored hairs. C. h. tadae was de- scribed by Tokuda and Kano (1936) as having a uniform gray dorsum with buffy brown and clove brown variations. None of these pelage colors ap- pears to be close to those of C. h. kurodai. C. h. ssp. from Fukien also has unicolored dorsal hairs, but they are Black. The ventral pelage of C. h. kurodai is Gray to Very Dark Gray. This condition is closest to C. h. indochinensis from Burma and C. h. watasei which have Grayish Brown and Dark Gray venters. C. h. tadae has a Dark Reddish Brown venter which was described by Tokuda and Kano (1936) as slightly paler than the dorsum and more silvery. The C. h. ssp. from Fukien has a Very Dark Grayish Brown venter. As with all C. horsfieldi, C. h. kurodai has bristles on the proximal third of the tail, a character noted by Tomes (1857) in the original de- scription of C. horsfieldi and mentioned both by Robinson and Kloss (1922) in the description of C. h. indochinensis and by Tokuda and Kano (1936) for C. h. tadae. The type-specimen of C. h. kurodai has a monocolor tail as does USNM 311473. The tail color is seen elsewhere only in the specimen of C. h. ssp. from Fukien. C. h. tadae (NSMT 4395) has a tail which is only slightly bi- colored, a characteristic observed in specimen USNM 332813 of C. h. kurodai. Tokuda and Kano (1936) reported that C. h. tadae has a tail so slightly bicolored that “. . . it cannot be said to be bicolored . . . .” All speci- mens from the other localities have bicolored tails. The vibrissae of C. h. kurodai as well as the specimens of C. h. watasei, C. h. indochinensis, and C. h. horsfieldi are unicolored, generally brown. VOLUME 90, NUMBER 3 463 However, the distal half of the vibrissae of C. h. tadae are white (Tokuda and Kano, 1936), a characteristic not noted in specimen NSMT 4395. The palatal length of C. h. kurodai is shorter than any of the other C. hors- fieldi. The length of the nasals is markedly shorter than in C. h. tadae and C. h. watasei but comparable to that of C. h. ssp. from Fukien, C. h. indo- chinensis from Tonkin and C. h. horsfieldi. The length of the upper tooth row is shorter than that of C. h. tadae, C. h. watasei, and C. h. indochinensis but comparable to that of C. h. horsfieldi and the Fukien specimen. The posterior edge of the palatine of C. h. kurodai is narrower than in any C. horsfieldi although it is close to that of C. h. horsfieldi and C. h. tadae. The height of the rostrum is comparable to all but C. h. tadae, which is not as high. In conclusion, the rostrum of C. h. kurodai is most similar in size to the C. h. ssp. from Fukien. It is shorter and somewhat more robust than that of C. h. tadae. Breadth of the braincase is less than in all but C. h. horsfieldi and C. h. ssp. from Fukien. The least interorbital breadth of C. h. kurodai is nar- rower than in C. h. tadae and C. h. horsfieldi, but close to that of other subspecies examined. C. h. kurodai is generally smaller than C. h. watasei, the largest C. hors- fieldi studied, and, is most similar to C. h. horsfieldi and C. h. ssp. from Fukien, although the latter is much darker. C. h. kurodai can be distin- guished from C. h. tadae by its generally smaller cranial measurements and grizzled Dark Reddish Brown (or grizzled Very Dark Brown) dorsal color in contrast to the solid, rustier Dark Reddish Brown of C. h. tadae: in addition, the rostrum of C. h. kurodai is shorter and more robust, and the ramus is relatively stout (Fig. 1A). Shou et al. (1966) described C. h. wuchihensis from Hainan. The de- scription suggests a typical C. horsfieldi but offers little with which we can compare our specimens. C. h. myoides from Kashmir was not examined. Remarks.—The relationships of C. h. kurodai appear rather clear except for C. h. tadae and C. h. ssp. from Fukien; both of the latter differ from C. h. kurodai but are represented by only one specimen each. The mainland specimens, C. h. indochinensis, exhibit considerable variation (Table 1) which indicates need for revision of this subspecies when more specimens are available. Five of the six named subspecies are insular, emphasizing the high degree of endemism among shrews (Repenning, 1967). Etymology.—This subspecies is named to honor Nagamichi Kuroda in recognition of his extensive studies of mammals of eastern Asia. Specimens examined.—Three, as follow: Taipei Hsien, Linkou (USNM 308115); Nantou Hsien, Lung Yuen Bridge 24°02’N, 121°08’E (USNM 332813); “Formosa” (not specific) (USNM 311473). 464 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Usary FSEE7L Fig. 1. A, Mandible, type of Crocidura horsfieldi kurodai n. ssp., (USNM 358115); B & C, Mandible, type of Crocidura russula hosletti n. ssp., (USNM 398640) labial and lingual views respectively; D & E, Mandible, Crocidura suaveolens (Pallas, 1811), (USNM 358672) lingual and labial views respectively; F—H, Mandible of Episoriculus fumidus (Thomas, 1913) lingual, posterior and labial views respectively. Crocidura horsfieldi tadae Tokuda and Kano Crocidura horsfieldi (Tomes, 1857) Jones, Lim and Cross, 1971:270; Jones, 1975:185. VOLUME 90, NUMBER 3 465 Crocidura tadae Tokuda and Kano, 1936 Tokuda and Kano, 1936:429; Kuroda, 1938:84, 1939:45, 1940:184; Okada, 1938:2: Chen, 1956:49; 1969:349. Crocidura horsfieldi tadae Tokuda and Kano, 1936 Ellerman and Morrison-Scott, 1951:76, 1966:76; Kuroda, 1952:286; Jones, Huang and Chang, 1969:49; Kuntz and Dien, 1970:33. Crocidura tadae was described on the basis of six specimens from Lan Yu (ie. Orchid Island or Kotéshé), southeast of Taiwan. Kuroda (1952) made C. tadae a subspecies of C. horsfieldi, and noted that it is not found on Taiwan proper. The habitat is described as farmyards and stone walls near houses (Tokuda and Kano, 1936). Measurements (from type-description)—Total length (5 specimens) mean = 95.6 mm (range 91-101); tail (5) 42.2 (39-45); hind foot (6) 12.2 (11.5-12.8); skull measurements (Table 1). Specimens examined—Lan Yu: Imourod 22°04’N, 121°32’E (NSMT 4395). Crocidura russula hosletti, new subspecies Crocidura horsfieldi (Tomes, 1857) Jameson, 1970:82. Holotype.—Adult male, skin and skull, USNM 398640, from 10 mi W Taichung, Taichung Hsien, Taiwan; obtained 11 October 1963 by E. W. Jameson, Jr., original no. EWJ 1150. Diagnosis—Generally small; tail short. Dorsal hairs tricolored, tips Dark Brown, middle portion lighter Dark Brown, and basal half very Dark Gray. Ventral hair of holotype Light Gray on throat and chest, with Dark Gray bases. Remainder of ventral pelage Grayish Brown with hairs unicolored to bases. Division of Light Gray and Grayish Brown portions rather abrupt, possibly a molt line. Tail only slightly bicolored; dorsum Dark Brown, venter lighter Dark Brown than dorsum; scales visible, although short hairs are rather numerous; bristles most abundant near base. Hind feet have Grayish Brown hairs dorsally. USNM 358671 has similar characteristics except that throat and chest lack Light Gray coloration. USNM 358670 has a Dark Reddish Brown dorsum which is darker than the other two skins. It also has Light Gray throat and chest and tail bristles like holotype, but 358671 has bristles on basal 21% of tail. Skins of both 358670 and 358671 are preserved in fluid. Skull small, condylo-incisive length short; mandible relatively short; height from condylar process to base short. Measurements.—Total length (3 specimens) mean = 108.7 mm (range 99-119); tail (3) 40.3 (39-42); hind foot (3) 13 (13); condylo-incisive length Hainan * Taken from type description (Shou et al., 1966). 466 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Table 1. Crocidura horsfieldi measurements, X and range (number of specimens measured). Hind Condylo- Total Tail foot incisive Palatal length length length length length _h. kurodai 104 44 12(2) —- 6.3 Taiwan 97-110(2) 42-46(2) 6.1-6.5(2) ._ h. tadae — — — 16.5(1) 6.6(1) Lan Yu uliaessp: 95(1) 35(1) Lit) 16.4(1) 6.5(1) Fukien . h. watasei 118 49.3 11.4 14 0 Ryukyus 112-123(3) 47-54(4) 11-12.2(4) 16.8-18.1(4) 6.6-7.5(4) . h. indochinensis —_ — — — 6.8(1) Tonkin . h. indochinensis 108 44.7 IIE 17.3 7.0 Burma 95-117(6) 42-47(6) 10-13(6) _—17.1-17.6(5) 6.8-7.2(6) . h. indochinensis 114 49 12.8 — = Thailand 103-125(2) 48-50(2) 12.5-13(2) . h. horsfieldi 116 49.9 Lee GM 6.5 Sri Lanka 111-120(10) 45-53(10) 10-15(11) 16.2-17.1(6) 6.4-6.7(9) . h. wuchihensis* — = 10.3 — — Hainan 10-10.5(2) Greatest Least Greatest breadth —interorbital maxillary Height Length UTR _braincase breadth width braincase . h. kurodai pal ten (ls) 3.7(2) Se 4.2(1) Taiwan 7.0-7.2(2) 5.1-5.3(2) _ h. tadae 7.4(1) 7.9(1) 3.9(1) 5.3(1) 4.1(1) anu SIs SID. HEED) TOD) 3.8(1) SJL) 4.0(1) Fukien | . h. watasei 7A 8.0 3.8 5.4 45 Ryukyus 7.1-7.9(4) 7.8-818(4) 35-41(4) 52°56) eeeeeaen . h. indochinensis Tesi) — 4.0(1) 5.2(1) — Tonkin . h. indochinensis U4! 8.1 4.0 De 45 Burma 7.2-7.6(6) 7.9-8.3(6) 3.9-4.1(6) 5.1-5.3(6) 4.4-4.6(4) . h. indochinensis Wed 8.1(2) --- 5.8 — Thailand 7.5-7.8(2) 5.7-5.9(2) . h. horsfieldi 7.0 35) 3.6 Dall 4.3 Sri Lanka 6.7-7.3(9) 7.2-7.7(5) 3.5-3.8(9) 4.9-5.3(9) 4.2-4.3(5) . h. wuchihensis -- 7.8(2) —_ — = VOLUME 90, NUMBER 3 Table 1. Continued. Length Height Mandibular LTR rostrum length C. h. kurodai Bye 1.8 —_ Taiwan 5.0-5.3(2) 1.7-1.9(2) C. h. tadae DE) Gls) 10.0(1) Iban) Yb Graliassp: 4.8(1) 1.8(1) @48)(b)) Fukien C. h. watasei HY 1.9 Wad Ryukyus 5.0-5.5(4) 1.8-2.1(4) 11.0-11.1(4) C. h. indochinensis 5.0(1) 2:0(1) 10.3(1) Tonkin C. h. indochinensis il 1.9 10.5 Burma 4,9-5.2(6) 1.8-2.0(6) 10.3-10.8(6) C. h. indochinensis = — = Thailand C. h. horsfieldi 4.9 1.8 9.9 Sri Lanka 4.6-5.0(10) 1.4-1.9(10) 9.5-10.5(10) 1.5-1.7(10) C. h. wuchihensis — = = Hainan Width posterior edge of Length palatine nasals C. h. kurodai Mell) 4.8(1) Taiwan C. h. tadae 2.3(1) 5.4(1) Lan Yu C. h. ssp 2.5(1) 4.9(1) Fukien C. h. watasei 5) 6.0 Ryukyus 2.4-2.5(2) 5.9-6.1(2) C. h. indochinensis 2.5(1) 4.9(1) Tonkin C. h. indochinensis — — Burma C. h. indochinensis — -- Thailand C. h. horsfieldi 22 4.8 Sri Lanka 2.1-2.3(6) 4.2-5.4(6) C. h. wuchihensis — — Hainan Length PM* 1.8 7182) SL) ILs7i(1L)) 1.6(4) Wea (i) od 1.6-1.7(6) 1.6 467 12 1.0-1.3(2) 1.0(1) Testi(ell) 1.3 1.2-1.4(4) e} 1 1.1-1.2(6) 1.2 1.0-1.3(10) 468 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Table 2. Crocidura russula measurements, x and range (number of specimens measured ). Condylo- Total Tail Hind foot incisive Palatal length length length length length C. r. hosletti 108.7 40.3 13(3) 17.8(1) al 99-119(3) 39-42(1) 6.9-7.1(4) C. r. quelpartis 114 45.8 13 18.9 1.2 98-131(8) 39-48(8) 12-15(8) 18.3-19.6(8) 7.0-7.3(8) C. r. chisai 123(1) 43(1) 15(1) 18.9(1) fell) C. r. rapax 118 47 (2) 13.5 Gs) 6.9 111-125 (2) 13-14(2) 6.8-6.9(2) C. r. vorax See 49 12.3 18.1 Tes: 108-125(3) 43-57(3) 12-13(3) 17.8-18.4(2) 7.0-7.7(3) C. r. pullata 129.8 45.4 1S 20 8.3 121-137(14) 40-50(14) 14-16(13) 19.4-20.4(8) 8.0-9.2(13) C. r. russula 106.5 40.8 APA 20.3 — 104-111(4) 38-43(4) 12-12.5(4) 19.5-20.6(4) Least Greatest Greatest Length _interorbital breadth maxillary Height UTR breadth braincase width braincase C. r. hosletti Cell 4.0 8.2(3) 5.6 4.6 7.5-7.9(4) 3.9-4.0(4) 5.6-5.7(4) 4.5-4.7(3) C. r. quelpartis Tool 4.0 8.3 Dell 4.5 7.5-8.0(8) 3.9-4.1(8) 8.0-8.6(8) 5.5-5.8(8) 4.2-4.7(8) C. r. chisai UBL) 4.0(1) 8.5(1) 5.7(1) 4.5(1) C. r. rapax 7.5 3.6 7.6(1) 5.2 4.5(1) 7.4-7.5(2) 3.5-3.7(2) 4.9-5.4(2) C. r. vorax 7.9 4.0 8.5 ll 4.6 7.4-8.0(3) 3.9-4.0(3) 8.4-8.5(2) 5.5-5.9(3) 4.5-4.7(2) C. r. pullata 8.9 4.4 9.2 6.2 4.6 8.5-9.3(13) 4.2-4.5(12) 8.9-9.7(10) 6.0-6.4(13) 4.5-4.7(6) C. r. russula 8.9 4.1 9.2 6.4 4.8 8.7-9.0(4) 4.0-4.3(4) 8.8-9.5(4) 6.3-6.5(4) 4.6-4.9(4) (1) 17.8; palatal length (3) 7.1 (6.9-7.1); greatest breadth braincase (3) 8.2 (8.2); length upper tooth row (3) 7.7 (7.5-7.9); greatest maxillary width (3) 5.6 (5.6-5.7); least interorbital width (3) 4.0 (3.9-4.0); height of braincase (3) 4.6 (4.5-4.7); length of lower tooth row (3) 5.2 (5.0-5.4); length of PM4 at cingulum (3) 1.8 (1.8); width of posterior edge of palate (2) 2.5 (2.5); Table 2. Continued. VOLUME 90, NUMBER 3 469 Width posterior Length Height Length Length edge of LTR rostrum PM* fe palatine . r. hosletti 5.2 LG) 1.8(4) 16) 2.4 5.0-5.4(4) 1.8-2.0(4) 1.3-1.4(4) 2.3-2.5(3) . r. quelpartis 5p) 1.8 1.8 el 742 5.1-5.4(8) 1.6-2.0(8) 1.7-1.8(8) 1.1-1.2(8) 2.2-2.4(8) . 1. chisai 5.3(1) ee) AG) eG) 2.5(1) 2 Mapax BUC) 1.9 1.8(2) UE) 2.4 1.8-1.9(2) 2.3-2.4(2) sit VOTAX 5.5 1.8 1.8(3) 1.2(3) 2.3 5.3-0.8(2) 1.8-1.9(3) 1.8-2.6(3) . r. pullata 6.1 1.9 2.0 1.3 2.8 5.9-6.3(14) 1.8-2.0(13) 2.0-2.1(14) 1.1-1.5(13) 2.0-3.0(13) . r. russula 6.1 1.9 2.0 1.3 2.8 6.0-6.2(4) 1.7-2.0(4) 1.9-2.0(4) 1.2-1.4(4) 2.7-2.9(4) Top of Infraorbital condylar foramen to Length Mandibular process anterior point infraorbital length to base premaxilla canal . rt. hosletti 10.9 3.8(4) 4.8 0.7 10.7-11.1(4) 4,6-5.0(4) 0.6—-0.9(4) . r. quelpartis 14! 4.2 4.9 0.6 11.1-11.8(8) 4,0-4.6(8) 4.7-5.1(8) 0.5-0.7(8) . 1. Chisai Te) 4.0(1) 4.7(1) 0.8(1) . T. rapax 10.6 3.9 4.4 0.7 10.5-10.6(2) 3.7-4.0(2) 4.3-4.5(2) 0.6-0.7(2) . t. vorax 11.3 4.0 4.6 0.9(2) 10.9-11.8(3) 3.9-4.0(2) 4,4-4.7(2) . r. pullata IPA) 4.5 5.5 0.8 12.2-12.9(13) 444.8(13) 5.45.9(13) 0.6-0.8(14) . r. russula 12.8 5.0 6.2 0.8 12,2-13.2(4) 4,7—-5.2(4) 5.8-6.5(4) 0.8—0.9( 4) height of rostrum (3) 1.9 (1.8-2.0); length of I1 at cingulum (3) 1.3 (1.3- 1.4); distance from infraorbital foramen to anterior point of premaxilla (3) 4.8 (4.6-5.0). Comparison.—Dorsal coloration of the holotype is the same as Allen 470 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON (1938) described for C. vorax (= C. russula vorax). However, USNM 358670 appears closest to C. suaveolens phaeopus (AMNH 56010, 56017, 56021, 56026, 56038). Allen (1938) described the venter of C. russula as like that of C. r. hosletti, whereas C. suaveolens had hairs of the venter “white-tipped,” “whitish-gray” and “white.” However, venters of the above specimens of C. suaveolens phaeopus in the AMNH are similar to C. r. hosletti. AMNH 56010 and 56026 have gray throats and chests as do the holotype of C. r. hosletti and USNM 358670. Tails of other subspecies of C. russula are more distinctly bicolored than C. r. hosletti, as are those of C. suaveolens. Distribution of tail bristles is variable in the subspecies of C. russula studied (our observations and pub- lished descriptions). The skull of C. r. hosletti is generally smaller than in other subspecies of C. russula (Table 2), although close to C. r. quelpartis, C. r. vorax and C. r. rapax. The palatal shape of C. r. hosletti is essentially the same as C. r. quelpartis, but the skull of the former is shorter and narrower and the braincase higher than in the latter. The braincase of C. r. hosletti is nar- rower than in C. r. vorax; that of C. r. rapax is narrower than in C. r. hosletti. The shape of the palate of C. r. vorax is similar to that of C. r. hosletti, but that of C. r. rapax is shorter and narrower than both. The mandible of C. r. hosletti (Fig. 1B and C) reflects the relationships de- scribed above, being larger than in C. r. rapax, and smaller than in C. r. vorax or C. r. quelpartis. Generally, C. r. hosletti is midway in size between C. r. vorax and C. r. rapax; C. r. hosletti is similar to C. r. quelpartis but somewhat smaller. The color of C. r. hosletti is closest to that described for the type of C. r. vorax (Allen, 1923) except that the tail is not distinctly bicolored. Allen (1923) described the ear of C. r. vorax as small and “less conspicuous than usual”; the ears of C. r. hosletti are relatively large and conspicuous. Biology.—C. r. hosletti was found (EWJ) to be common in cultivated fields in lowlands of Taichung Hsien, where it apparently outnumbers C. attenuata. It was reproductively active in February when 3 gravid fe- males were collected. Two had 3 embryos and the other had 4 embryos. None was nursing. Etymology.—This subspecies is named to honor our friend, the late Professor Sherman A. Hoslett. Specimens examined.—Taipei Hsien, Tamsui 25°10’N, 121°26’E (USNM 398670). Taichung Hsien: 10 mi W Taichung 24°10’N, 120°38’E (USNM 398640). Taiwan (Not specific) (USNM 358671). Crocidura suaveolens (Pallas, 1811) This is the first report of C. suaveolens from Taiwan. The only known specimen is a skull (USNM 358672) collected in a bamboo forest, near VOLUME 90, NUMBER 3 471 Table 3. Crocidura suaveolens measurements, and range (number of specimens measured ). Palatal Length length UTR C. suaveolens 6.9(1) 7.4(1) Taiwan C. s. shantungensis — — C. s. phaeopus 6.6 TP 6.3-7.1(16) 7.0-7.7(14) C. s. ilensis 7 O@D) 7.4(1) C. s. lignicolor SalLCL) 8.5(1) Height Mandibular rostrum length C. suaveolens ILL) 10.5(1) Taiwan C.s. shantungensis —_1.7(1) — C.s. phaeopus 16 9.8 1.5-1.8(30) 9.7-10.2(10) C.s. ilensis E3710) —_ C.s. lignicolor 2:0(1) _ Top of condylar process to base C. suaveolens Bec (1) Taiwan C. s. shantungensis 3.9( 1) C. s. phaeopus 3.6 3.2-4.1(20) C. s. ilensis 3.8(1) C. s. lignicolor 4.3(1) Feng ChiHu, 1,440 m. The cranial Least Greatest interorbital maxillary Length breadth width TAR 30 CG) 5.0(1) 5.3(1) — 5.3(1) 5.3(1) 3.6 5.0 5.0 3.4-3.8(17) 4.8-5.1(17) 4.7-5.4(20) 3.91) 5.1(1) BA) 4.5(1) Gy) 6.1(1) Width posterior Length Length edge of PM* Ie palatine ed (CL) ILBGD) 2.4(1) ES Gy) — DFAS) led JbeJL Yt 1.6-2.1(20) 1.0-1.3(20) 2.1-2.9(17) ea) ee) Da) 1.9(1) 1.2(1) 1.9(1) Infraorbital foramen to Anterior Infraorbital point of canal premax length 4.8(1) 0.7(1) 4.3(1) 0.5(1) 4.3 0.6 3.8—5.0(19) 0.5-0.9( 19) 4.2(1) 0.6( 1) Orek) 0.8( 1) and mandibular measurements (Table 3) are close to but slightly smaller than those of C. russula hosletti. Massiveness of the ramus is the most useful character separating C. suaveolens and C. russula (Fig. 1D and E). As stated by Ognev (1962), the lower jaw of C. suaveolens is “much weaker, more slender and smaller than in...” C. russula. 472 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Generally, the specimen of C. suaveolens from Taiwan is larger than C. s. phaeopus, C. s. shantungensis and C. s. ilensis and smaller than C. s. lignicolor (Table 3). C. suaveolens from Taiwan is larger than its geo- graphically closest relatives, C. s. phaeopus and C. s. shantungensis, in the following characters: greatest maxillary width, height rostrum, length of rostrum from anterior edge of infraorbital canal to anterior tip of pre- maxilla and mandibular length. Discovery of this species and C. russula on Taiwan adds two more Pale- arctic species to the island fauna. More intensive collecting for small shrews on the island may yield additional material of C. suaveolens on which a more comprehensive description can be based. Measurements.—Table 3. Specimens examined.—Chiai Hsien: Feng Chi Hu 23°30’N, 120°41’E (USNM 358672). Suncus murinus (Linnaeus) Sorex murinus Linnaeus, 1766 Swinhoe, 1870:620. Crocidura murina (Linnaeus, 1766) Aoki, 1913:272, 1914:30; Horikawa, 1925:224. Suncus murinus (Linnaeus, 1766) Horikawa, 1925:233; Kuroda, 1925:2; Chen, 1956:48, 1969:349; Jones, Huang, Chang, 1969:49; Kundin et al., 1972:270; Kuntz and Dien, 1970: 33; Jones, Lim and Cross, 1971:269; Jones, 1975:185. Crocidura (Pachyura) murina (Linnaeus, 1766) Aoki, 1930:13, 1933:79; Tanaka, 1936:311; Tateishi, 1938:516. Suncus murinus murinus (Linnaeus, 1766) Ellerman and Morrison-Scott, 1951:65, 1966:64. Sorex myosurus Pallas, 1785 Swinhoe, 1864:382. Crocidura caerulea (Shaw, 1800) Horikawa, 1925:224. Suncus caeruleus (Shaw, 1800) Kuroda, 1925:1; Harrison and Audy, 1951:179. Suncus murinus swinhoei (Blyth, 1859) Kuroda, 1952:285. Pachyura swinhoei (Blyth, 1859) Gee, 1929-1930:52. Crocidura murina swinhoei (Blyth, 1859) Horikawa, 1932b:249. Crocidura (Pachyura) murina swinhoei (Blyth, 1859) Takashima, 1930:199; Horikawa, 1932a:35. Suncus myosurus swinhoei (Blyth, 1859) Okada, 1938:2; Kuroda, 1938:81, 1940:177. VOLUME 90, NUMBER 3 473 Crocidura muschata Hatori, 1915 Hatori, 1915:57, 1919:234. Suncus murinus commonly occurs in houses and other buildings as well as open city sewers, livestock pens, etc. It is less common in cultivated areas near houses. We found it only below 300 m. McNeill et al. (1968) found this species living in the coral rock walls near the villages of MaKung, Peng Hu Hsien (Pescadores). Mao (1970) reported that the habu viper (Trimeresurus mucrosquamatus) is a predator of Suncus murinus. Measurements.—Total length males (7 specimens) mean = 211.9 mm (191-223), females (4) 191.3 (184-198); tail males (7) 80.3 (70-89), females (4) 71.3 (67-78); hind foot males (7) 21.6 (20-23), females (4) 20.3 (20-21). Specimens examined.—Chang Hua Hsien: Hua-Tan 24°02’N, 120°32’E (USNM_ 313633, 313634, 313635); Chi Hu 23°58’N, 120°28’7E (USNM 332818); Pu Yen 24°00’N, 120°28’E (USNM 313650); Shan Sheng (?co- ordinates) (USNM 313640, 313641, 313646, 313647, 313648, 313649). Hsin Chu Hsien: Pei Pu 24°42’N, 121°03’E (USNM 313643, 313644). Hualien Hsien: 3 mi SW Hualien 23°58’N, 121°36’E (USNM 330006, 330007, 330008, 330009). Kaohsiung Hsien: Tsoying 22°41’N, 120°17’7E (USNM 294134, 294135, 356347, 356348); Mei Nung 22°54’N, 120°32’E (USNM 332819). Peng Hu Hsien: Hu Hsi 23°35’N, 119°390’E (USNM 294136, 294137, 294138); Lin Tou 23°34’N, 119°38’E (USNM 332820, 332821, 332822); Pachnau 23°22’N, 119°29"E (USNM 294728, 294542, 294543): Gyoo-To 23°36’N, 119°30’°E (USNM 294559, 294560, 294561, 294562, 294563); (no specific locality) (USNM 294557, 294558, 294564, 294565, 294566, 294567, 330016, 330017, 330018, 330019, 332823, 332824). Ping Tung Hsien: Chao Chow 22°33’N, 120°32’E (USNM 297862, 299956, 313637, 332816, 332817, 330001, 356350); San Hsing 22°33’N, 120°33’E (USNM 356349). Taichung Hsien: Ma-an Liao, 6 mi S. Tung Hsi 24°15’N, 120°49’E (USNM 294130); Taichung 24°09’N, 120°41’E (USNM 294131); 4 mi SE Taichung (USNM 294132, 294133). Taipei Hsien: Ali Lao 25°17’N, 121°36"E (USNM 330004, 330005); Pei Tou 25°08’N, 121°29’E (USNM 313638, 313639); Shih Lin 25°05’N, 121°31’E (USNM 313645); Tien Mou 25°05’N, 121°31’E (USNM 330010, 330011, 330012, 330013, 330014, 330015); Taipei 25°03’N, 121°30’E (USNM 305926, 305927, 305928, 305929, 305930, 305931, 305932, 305933, 308418, 308419, 308420, 308421, 308422, 308520, 313631, 313632, 313636, 313642, 313651, 330002, 330003, 332815, 358116, 308129). Tamsui 25°10’N, 121°26’E (USNM 358117, 358118, 358119, 358120, 308121, 358122, 358123, 358124, 358125, 358126, 358127, 358128, 358130). Literature records—Chiai Hsien: 23°29’N, 120°27’E (Kuroda, 1925, 1938). Peng-Hu Hsien: Makung 23°34’N, 119°34’E (McNeill et al., 1968). Tainan Hsien: Tainan 23°00’N, 120°11’E (Kuroda, 1938). Taipei Hsien: Yangmingshan 25°09’N, 121°33’E (McNeill et al., 1968). Miao-Li Hsien: A474 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Chu-Nan 24°41’N, 120°S27E (Kundin et al.) 1970).)\ Hao M@onmsbisien (Mao, 1970). Episoriculus fumidus (Thomas) Soriculus caudatus (Horsfield, 1851) Jones; im) and Cross, 1971-269: Soriculus fumidus Thomas, 1913 Thomas, 1913:216; Aoki, 1913:270, 1914:30; Horikawa, 1925:224, 1932a: 13, 1932b:249; Cabrera, 1925:125; Tanaka, 1936:311; Kuroda, 1938:80, 1940:174; Okada, 1938:3; Chen, 1948:41, 1956:47, 1969:349; Kuntz and Dien, 1970:33. Soriculus caudatus fumidus Thomas, 1913 Ellerman and Morrison-Scott, 1951:59, 1966:59; Kuroda, 1952:285; Jones, Huang and Chang, 1969:48. Episoriculus fumidus (Thomas, 1913) Jameson, 1970:80; Jones, 1975:184. Chodsigoa sodalis Thomas, 1913 Thomas, 1913:217; Aoki, 1913:270, 1914:30; Horikawa, 1925:224, 1932a: 13, 1932b:249; Cabrera, 1925:127; Kuroda, 1938:80; Okada, 1938:3; Chen, 1948:42, 1956:48, 1969:349; Ellerman and Morrison-Scott, 1951: 61, 1966:61; Jones, Huang and Chang, 1969:48; Jones, 1975:184. Thomas (1913) described this shrew from a series of skins and skulls from Alishan, Chiai Hsien, placing it in the genus Soriculus. Is is common and widely distributed in the mountains. Taken primarily in hardwood and coniferous forests above 1,000 m, it has also been collected in dwart bamboo on Ho Huan Shan, 3,200 m. Although many specimens were examined, no gravid females were found. Ellerman and Morrison-Scott (1951) established the subgenus Episori- culus on the basis of size, proportions, and certain external features; they included the species S. caudatus and S. leucops, but questioned the place- ment of fumidus. On the basis of mandibular and dental characters, Repen- ning (1967) elevated Episoriculus to generic level and illustrated some im- portant cranial features of Episoriculus caudatus. Kuroda (1925) intimated that E. fumidus might be a subspecies of E. caudatus, but a comparison of our specimens with Repenning’s (1967) illustration shows that the mandible of E. fumidus differs from that of E. caudatus in having the angular process quite long and curved and a high coronoid process with converging sides (Fig. 1F and H). The lower incisor is long and in this respect E. fumidus resembles Neomys. Thus E. fumidus is clearly different from E. caudatus and in most respects fits comfortably into Repenning’s (1967) description of Episoriculus. VOLUME 90, NUMBER 3 AT5 Along with the original series of E. fumidus was a skull (without an accompanying skin) which Thomas (1913) described as the holotype of Chodsigoa sodalis. This skull was placed in Chodsigoa because it lacked the fourth upper unicuspid which is characteristic of Soriculus (and now Episoriculus). Thomas (1913) closed the description of Chodsigoa sodalis with the following comment. “Its external appearance is probably not very dissimilar from that of the other Arizan shrew, Soriculus fumidus, as Mr. Goodfellow brought home the typical skull as being just an extra skull of that species.” In many efforts at Alishan and other seemingly appropriate sites, we have failed to find Chodsigoa sodalis, and have concluded that C. sodalis was based on an aberrant specimen of E. fumidus. Dr. G. A. Corbet (BMNH) has examined the types of both species and informs us that they seem indistinguishable except for the rudimentary unicuspid and a difference in age. Measurements.—Total length males (8 specimens) mean = 112.1 mm (range = 105-126), females (9) 111.1 (103-128); tail males (8) 47.0 (45-50), females (9) 48.0 (44-52); hind foot males (7) 13.1 (12-14), females (9) 12.8 (11-14); condylo-incisive length (1) 18.1; greatest breadth braincase (2) 9.5 (9.5); length upper tooth row (2) 7.8 (7.5-8.0). Specimens examined.—Chiai Hsien: Alishan 23°32’N, 120°48’E (USNM 261036, 332804, 332805, 332806, 332807, 332808, 332809, 332810, 358108); 0.1 km E Alishan Station (USNM 358108); 0.2 km E Alishan Station (USNM 358657); Wu-feng (2,500 m), 10 km SW Alishan (EWJ 1158) 2 km W Alishan (USNM 358658). Nantou Hsien: Wushe 24°02’N, 121°08’E (USNM 332811, 332812), Yin Feng, Wushe (USNM 332802, 332803): Chuei Feng 24°05’N, 121°11’E (USNM 332800, 332801); Ho Huan Shan 24°09’N, 121°16’E (USNM 358659, 358580). Literature records——Nantou Hsien: Lung Yuen, 5 km SW Wushe (Philipps, 1966); 24 km E Wushe (Jameson, 1970), 32 km NE Wushe (Jameson, 1970). Taichung Hsien: Lishan 24°15’N, 121°15’E (Jameson, 1970). Chimarrogale himalayica (Gray, 1842) Chimarrogale himalayica was first reported and fully described from Taiwan by Jones and Mumford (1971). Four specimens were collected in boulder strewn mountain streams near the villages of Pei Chang and Meichi, Nantou Hsien. Measurements.—Total length males (2 specimens) mean = 122.5 mm (range = 115-130), females (2) 110 (109-110); tail males (2) 85 (80-90), females (2) 96.5 (92-101); hind foot males (2) 26 (25-26), females (2) 23 (23); condylo-incisive length males (2) 25.0 (24.9-25), females (2) 24.8 (24.6-25.0); length upper tooth row males (2) 11.2 (11.0-11.3), females 476 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON (2) 11.0 (10.9-11.1); greatest breadth braincase males (2) 12.9 (12.9), fe- males (2) 12.9 (12.8-13.0); greatest maxillary width males (2) 7.9 (7.8- 7.9), females (2) 7.6 (7.5-7.6); height braincase males (2) 6.7 (6.6-6.7), fe- males (2) 6.8 (6.7-6.8). Specimens examined.—Nantou Hsien: Pei Chang Chi (stream) 24°02’N, 120°5V’E (RMNH 20963); Meichi (stream) 23°59’N, 120°54’7E (USNM 358139, 358140, 358141). Anourosorex squamipes yamashinai Kuroda Anourosorex squamipes Milne-Edwards, 1872 Jones, Lim and Cross, 1971:269; Jones, 1975:186. Anourosorex squamipes yamashinai Kuroda, 1935 Kuroda, 1935:288, 1938:85, 1940:187, 1952:286; Tanaka, 1936:313; Okada, 1938:3; Chen, 1948:46, 1956:49, 1969:349; Ellerman and Morri- son-Scott, 1951:87, 1966:87; Jones, Huang and Chang, 1969:50; Kuntz and Dien, 1970:33; Jameson, 1970:82. This shrew was first found in Taiwan by Kuroda (1935), who described it as an endemic subspecies. It apparently occurs throughout the forested mountains from about 300 m (the subtropical zone of Kuroda, 1952) to above timberline at elevations of more than 3,000 m (subarctic zone of Kuroda, 1952). This shrew is most abundant in hardwood forests between 1,500 and 2,500 m (upper temperate zone of Kuroda, 1952). It has also been captured in a cornfield near Wushe and in streamside secondary growth near Meichi. At the upper limits of its distribution it inhabits bam- boo-covered slopes where it was captured in surface runways of Microtus kikuchii. There the bamboo seldom exceeds 30 cm in height. Measurements.—Total length males (13 specimens) mean = 105.2 mm (range = 95-113 mm), females (10) 99.6 (90-109); tail males (13) 10.2 (7-11), females (10) 9.8 (8-12); hind foot males (13) 15.3 (13-18), females (10) 14.9 (10-16.6); condylo-incisive length males (4) 25.2 (24.6-26.2), fe- males (2) 25 (24.3-25.7); palatal length males (5) 10.6 (10.1-11.3), females (2) 11.1 (10.5-11.6); greatest breadth braincase males (5) 13.3 (12.3-13.8), females (1) 13.2; length upper tooth row males (6) 11.2 (10.6-12), females (2) 11.3 (11.1-11.4). Specimens examined.—Chiai Hsien: Alishan 23°32’N, 120°48’E (USNM 261040, 261041, 261042, 261043, 261044, 261045, 261046, 261047, 332828, 332829); 0.1 km E Alishan Station (USNM 358132, 358133, 358134); 0.2 km E Alishan Station (USNM 358135); Feng Chi Hu 23°30’N, 120°41’E (USNM 398136, 358137). Nantou Hsien: Wushe 24°02’N, 121°08’E (USNM 332825, 332826, 332830); 4.1 km W and 2.7 km S Wushe (USNM 358131); Chuei feng 24°05'N, 121°11V’E (USNM 332831); Meichi 6 km W Wushe 23°59’N, 120°54’E (USNM 358138); Sung Kong forest (?coordinates) (BPBM 171). VOLUME 90, NUMBER 3 477 Literature records.—Ilan Hsien: Taiheizan (presently known as Tai- pingshan) 24°30’N, 121°38’E (Kuroda, 1935). Nantou Hsien: 10 mi E Wushe (Jameson, 1966). lA, iiloy 2a. 2b. 3a. 3b. Aa. Ab. ae ob. 6a. 6b. Uo 7b. 8a. 8b. la. lb. 2a. 2b. 3a. 3b. Aa. Ab. Da. Ob. 6a. 6b. Ta. Tb. Key to Skulls of the Soricidae of Taiwan Seven teeth in upper tooth row (= 14) Anourosorex squamipes yamashinai More than seven teeth in upper jaw 2a Nine teeth in upper jaw (= 18) Suncus murinus Eight teeth in upper jaw (= 16) 3a Tips of teeth pigmented, chestnut Episoriculus fumidus Tips of all teeth white 4a Skull more than 24 mm long Chimarrogale himalayica Skull less than 23 mm long Crocidura oa Upper tooth row longer than 8.5 mm _ Crocidura attenuata tanakae Upper tooth row shorter than 8.0 mm 6a Greatest maxillary width less than 5.4 mm 7a Greatest maxillary width more than 5.4 mm Sa Occurring on main island of Taiwan Crocidura horsfieldi kurodai Occurring on Lan Yu Crocidura horsfieldi tadae Mandibular length measuring 10.5 mm or less Crocidura suaveolens Mandibular length exceeding 10.6 mm Crocidura russula hosletti Key to Skins of the Soricidae of Taiwan Tail shorter than hind foot Anourosorex squamipes yamashinae Tail longer than hind foot 2a Edges of feet and tail with fringes of stiff hairs Chimarrogale himalayica Edges of feet and tail without fringes of stiff hairs 3a Tail with long bristles 4a Tail without long bristles Episoriculus fumidus Hind foot longer than 20 mm Suncus murinus Hind foot shorter than 15 mm_ Crocidura 5a Tail bristles limited to proximal third of tail Crocidura horsfieldi 6a Tail bristles occurring over more than proximal third of tail 7a Occurring on main island of Taiwan Crocidura horsfieldi kurodai Occurring on Lan Yu Crocidura horsfieldi tadae Tail longer than 45 mm; total length greater than 120 mm Crocidura attenuata tanakae Tail shorter than 45 mm; total length less than 120 mm Crocidura russula kurodai 478 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Discussion Taiwan lies off the coast of the Chinese mainland where the Palaearctic and Oriental Regions interdigitate over a broad front. The lowlands of Taiwan are subtropical and the shrews, Crocidura and Suncus, inhabit- ing this climatic region are widespread in the Old World. Mountains rise sharply from near the eastern seacoast to treeless alpine slopes of dwarf bamboo at 3,200 m and more. Four vegetative zones occur in the moun- tains, from the plains which are extensive on the western part of the island to the high ridges on the east (Tsukada, 1966); shrews occur over a wide range of elevations. Chimarrogale himalayica is contined to lower ele- vations in the mountains presumably by virtue of being restricted to streams of moderate speed that flow through the narrow mountain valleys. Anourosorex squamipes occurs from warm temperate forests of Cunning- hamia konishii, Podocarpus macrophyllus and Cephalotaxus wilsonia with a continuous distribution to above the boreal forests of Tsuga chinensis, Abies kawakamii and Picea morrizonicula. Episoriculus fumidus lives in forests, open grasslands, and dwarf bamboo from approximately 1,800 m to at least 2,300 m. Crocidura attenuata, as noted above, occurs in the sub- tropical lowlands, but also ranges to 1,225 m. Similarly, Crocidura hors- fieldi kurodai ranges from about sea level to above 1,200 m. The little known Crocidura russula and Crocidura suaveolens are only known to occur in the lowlands and mountains, respectively, and both are Palearctic in distribution. Under the cooling effects of the Pleistocene, the vegetative zones ex- tended into lower elevations. During the Tali Glacial Age (equivalent to the early Wisconsin period of North America), the boreal coniferous forests, which now occur above 2,400 m, were at 750 m (Tsukada, 1966). This sug- gests a drop of 8° to 11°C in the mean annual temperatures (Tsukada, 1966). During this period, Taiwan was connected to the mainland (Tsukada, 1966) at an estimated “glacial low stand” of —85 to -130 m (McIntyre et al., 1976); the Taiwan Strait is less than 100 m deep. Clearly, the climate was such that A. squamipes, the various species of Crocidura, and Chi- marrogale himalayica could have migrated from the mainland to Taiwan, if they had not already existed on the island. Anourosorex occurred far to the north at this time and is known from the middle Pleistocene of Honshu, Japan (Shikama and Hasegawa, 1958). Tsukada (1966) noted at least two subsequent cool periods on Taiwan; the last, between ca 25,000 and 14,000 BP was characterized by a mean temperature drop from 2° to 6°C. Later (14,000 to 12,000 BP) there was a rise in the mean annual temperature to about 2° to 3°C above recent levels. One must pre- sume that the drastic lowering of temperature during the Tali effected a total departure of Suncus from Taiwan, if it in fact occurred there prior to that time. VOLUME 90, NUMBER 3 479 This suggests that the antiquity of the Taiwan shrews is greater than 14,000 BP. Episoriculus fumidus is presumably the oldest soricid resident on Taiwan and is the only species known to be endemic. Anourosorex squamipes, Crocidura horsfieldi, C. russula and C. attenuata, judging from their present altitudinal distribution, could have survived the Tali Age on Taiwan. Gene flow during the period of the land bridge would account for their affinity with mainland forms. Chimarrogale himalayica could have conceivably followed the same pattern, for its present known distribution extends to 750 m and its apparent close relationships with C. himalayica leander suggest a migration during the land connection during the Tali Age. The fact that S. murinus is always found in close rela- tion with man’s abodes lends credence to the theory that it entered with man; this association with man throughout southern Asia suggests the probability that there have been repeated introductions on Taiwan via cargo or human immigration. This sequence of events could account for the low degree of endemicity among the shrews of Taiwan. Acknowledgments We thank the following individuals and their institutions for loaning comparative material used in our study: Dr. Sydney Anderson, AMNH., Dr. G. B. Corbet and Ms. P. D. Jenkins, BM(NH); (the late) Mr. Kitti Thonglongya, Applied Scientific Research Corp., Bangkok, Thailand; Drs. Y. Imaizumi and M. Yoshiyuki, NSMT; Dr. Luis de la Torre, FMNH; Dr. A. M. Husson, RMNH and Dr. Henry Setzer, USNM. We thank Drs. G. B. Corbet, David H. Johnson, and Charles A. Repenning for valuable suggestions during the study; Drs. John H. Cross and R. H. Watten, U.S. Naval Medical Research Unit #2, Taipei, for supporting parts of the field work, and Diana B. Jones for typing manuscript. Literature Cited Allen, G. M. 1923. New Chinese insectivores. Amer. Mus. Novitat. 100:11 pp. 1938. Mammals of China and Mongolia. Natural History of Central Asia. Amer. Mus. Nat. Hist. 11(1):620 pp. Aoki, B. 1913. A handlist of Japanese and Formosan mammals. Annot. Zool. Jap. 8(2):261-353. ———_—.. 1914. List of Taiwan mammals. Jour. Nat. Hist. Soc. Taiwan 4(14—19): 30-33. ————, 1930. The mammal fauna of Formosa and its surroundings. Chigaku Zasshi (Jour. Geogr.) 42:501-509. (In Japanese) 1933. Explanations on the Formosan mammals. Zool. Mag. 45(532-533): 79-80. (In Japanese) Cabrera, A. 1925. Genera mammalium. Insectivora and Galeopithecia. Mus. Nac. Ciencas Nat. 232 pp. 480 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Chen, J. T. F. 1948. g& y s I i \ by shelflike projection of sternal region, giving “half-moon” effect. B, Parthenope Vi pro] § § § i (Platylambrus) serrata, west of Sanibel Island, Florida. Note fully exposed gonopores on sternal plate, giving rounded effect. Scales equal 5 mm. 518 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Discussion Parthenope serrata and Parthenope granulata are completely rede- scribed in another paper (Gore and Scotto, in press). The synopsis of fea- tures presented here may be used to separate adults of the two species, but complete descriptions should be consulted for positive identification, espe- cially of younger adults or juveniles. The gonopods of the males (Fig. 1) and the gonopores of the females (Fig. 2) allow the species to be easily separated. In P. granulata the teeth on the outer margin of the hand are nearly straight, acutely tri- angular or even lanceolated and constricted somewhat at their bases, and have tips that project directly outward or often obliquely forward. In P. serrata these same teeth are usually more widely and regularly triangular and spinous, with their tips usually curved and directed more toward the fingers. In P. granulata, the posterolateral spine of the carapace has its posterior margin convex or oblique, not transverse, the tip is more blunt and often somewhat foliate, pointing upward and outward, and the entire spine appears to be directed obliquely posteriad. In P. serrata the pos- terior margin of this spine is transversely straight or very slightly curved, the tip is usually distinctly acuminate, and the entire spine is directed nearly straight outward or only very slightly obliquely posteriad; some adult specimens show this latter feature more noticeably than others. Parthenope serrata is compressed dorsoventrally, but the carapace is more inflated than P. granulata, the tuberculosity more diffusely dispersed dorsally, the granules and tubercles smaller and more numerous. In P. granulata the carapace is more compressed, and appears more angular dorsally (although this distinction is often not easily discerned in sparse material), the tuberculosity is more distinct and noticeable, the granules larger, more widely separated, and more regularly arranged, especially on the branchial ridges and interbranchial depressions. The posteriorly directed spines on the intestinal and posterior margins are, in many cases, more protuberant in P. granulata while appearing more blunt or reduced in P. serrata. When they are developed in the latter species they often curve distinctly upward. There usually appears a series of 4 or 5 clearly demarked granules in a more or less transverse row just above the in- testinal tubercle, and extending toward the two posterior tubercles in P. granulata; these granules are indistinct or lacking in P. serrata. The carapace of the latter species is often more pilose than the former so that much of the carapacial features appear indistinct. The pilosity on the cheliped marginal teeth is variably heavy in both species. These features are all easily compared in Plates 4 and 5. Sexual dimorphism in carapace shape appears in both species with males being more rounded anterolaterally, with a narrower carapace, while fe- VOLUME C0, NUMBER 3 519 Plate 4. Parthenope (Platylambrus) serrata (H. Milne Edwards, 1834). Male, off Honduras, Central America, R/V John Elliott Pillsbury station P-1361. A, Dorsal view: B, Ventral view. Note pilosity on margins of carapace and chelipeds. Parthenope (Platy- lambrus) granulata (Kingsley, 1879). Male, off central eastern Florida coast, SIFP 89: 1154. C, Dorsal view; D, Ventral view. Scales equal 10 mm. males appear more angular anterolaterally and have a noticeably wider carapace. Decided differences appear in the gonopores of the females of the two species. In P. granulata, the gonopore is raised up slightly from the plane of the sternal plate, directed more anteriorly, sheltered under a shelflike ridge, and appears distinctly half-moon shaped (Fig. 2A). In P. serrata, the mature female gonopore is rather flattened to the plane of the sternal plate, appears directed obliquely upward, is distinctly exposed without a shelflike ridge, and noticeably rounded (Fig. 2B). In P. serrata the abdom- inal somites are only slightly carinate transversely, the seventh somite tip is more acutely angular, and the outer margins sweep downward more abruptly to somite 6 than they do in P. granulata. In the latter species 520 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Plate 5. Parthenope (Platylambrus) serrata (H. Milne Edwards, 1834). Female, off Sanibel Island, Florida, SIFP 89:2526. A, Dorsal view; B, Ventral view. Parthenope (Platylambrus) granulata (Kingsley, 1879). Female, off Cape Canaveral, Florida, SIFP 89:1438. C, Dorsal view; D, Ventral view. Scales equal 10 mm. the abdominal somites are often sharply carinate transversely, somite 7 is more rounded and expanded and the tip is not nearly as acute (com- pare Plates 5B, D). Revised Synonymies for Parthenope (Platylambrus) serrata and P. (P.) granulata It has not been possible to examine every specimen noted in the litera- ture as Parthenope serrata to determine if it was correctly identified. How- ever, based on descriptions, and more importantly, illustrations, a partly revised synonymy for the two species has been constructed. Under Parthen- ope incertae sedis are placed literature citations consisting of incomplete descriptions, mere species listings, or those which have not yet been re- examined for this study. VOLUME 90, NUMBER 3 521 Parthenope (Platylambrus) serrata (H. Milne Edwards, 1834) (Plates 1, 4A, B; 5A, B; Figs. 1E-H, 2B) Lambrus serratus H. Milne Edwards, 1834:357.—Holthuis, 1959:191.— [Not White, 1847:12 (= Lambrus hoplonotus var. granulosus Miers, 1879) ]. Lambrus lupoides White, 1847:12 [Nomen nudum; specimen c, as re- stricted herein ]. Lambrus crenulatus Saussure, 1858:429 [13], pl. 1, figs. 4, 4a—Gundlach and Torralbas, 1899:301, text-fig. on p. 303 [In reprint, 1917:12, pl. [2], fig. 5 (Not seen) ]. Lambrus melanodactylus Desbonne, in Schramm, 1867:21 [Manuscript name]. Platylambrus serratus—A. Milne Edwards, 1878:156, pl. 30, figs. 1-Ic. [Illustration based on lectotype selected herein].—Rathbun, 1901:80. Lambrus (Lambrus) serratus—Young, 1900:102 [Key], 105. Parthenope (Platylambrus) serrata-—Rathbun, 1919:346; 1925:516 [In part], pls. 180, 181, and pl. 275, figs. 7-10 (After A. Milne Edwards, 1878; lectotype by present designation).—Boone, 1930:117 [In part], pl. 36, figs. A, B.—Chace, 1956:162.—Bullis and Thompson, 1965:13 [In part, R/V Oregon stations, and R/V Silver Bay stn. 71 only|.—Leary, 1967: text-fig. (p. 45), 50 [Listed].—Yang, 1971:166, figs. 1-9 [Larval development].—Felder, 1973:45 [Key], pl. 6, fig. 8 [Right chela].— Zeiller, 1974:100, color plate. Lambrus (Platylambrus) serratus.—Flipse, 1931:93. Type-Locality —“L’Océan indien” by original designation; erroneous lo- cality for the West Indies. Veracruz, Mexico by present selection of male lectotype; latter in Muséum National d'Histoire Naturelle, Paris, France. Parthenope (Platylambrus) granulata Kingsley, 1879 (Plates 3A, 4C, D; 5C, D; Figs. 1A-D, 2A) Lambrus granulatus Kingsley, 1879:150 [As noted, female syntype, USNM 55696; lectotype as selected herein]. Platylambrus serratus —Aurivillius, 1889:59, pl. 4, fig. 8—Hay and Shore, HONG: AG3 pl. 39), ie. 7. Parthenope (Platylambrus) crenulata.—Verrill, 1908:417, pl. 28, fig. 5 [Not pl. 27, as in text]. Parthenope crenulata—Verrill, 1922:155, text-fig. 12. Parthenope (Platylambrus) serrata—Rathbun, 1925:516 [in part*, not pls. 180, 181, and pl. 275, figs. 7-10 (all = P. serrata)|—Boone, 1930: 117 [In part, not pl. 36, figs, A, B (= P. serrata)|.—Springer and Bullis, 522 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 1956:22 [Listed].—Bullis and Thompson, 1965:13 [In part, R/V Silver Bay, stn 54, R/V Combat, stn 397, only].—Williams, 1965:267, text-figs. 247, 252B. T ype-Locality —Tortugas, Florida, 9 fathoms. Lectotype by present selection in National Museum of Natural History, Washington, D.C., USNM_ 55696. Parthenope (Platylambrus) incertae sedis Lambrus crenulatus.—Stimpson, 1860:201; 1871a:129; 1871b:101 [Discus- sion|].—Martens, 1872:85. Platylambrus serratus—Kingsley, 1880:390.—A. Milne Edwards, 1880:5.— Smith, 1886:629 [25].—Rathbun, 1897:12; 1898:261; 1900:514 [Key].— A. Milne Edwards and Bouvier, 1923:355.—Balss, 1924:181.—Flipse, 1930:86 [Listed]. Lambrus serratus—Miers, 1886:94 [Listed], 97.—Ortmann, 1893:415.— Moreira, 1901:62, 129, 130 [Synonymy].—Flipse, 1930:84 [Listed]. Parthenope (Platylambrus) serrata—Boone, 1927:42.—Rathbun, 1933:38 [Key], 39; 1935:114 [Discussion]—Chace, 1956; 162 [In part, MHN LaSalle material only ].—Righi, 1966:140.—Rodrigues da Costa, 1968:143; 1969:177 [Abstract].—Turkay, 1968:251.—Godcharles and Jaap, 1973: 48.—Coélho and Ramos, 1972:205 [Listed].—Pequegnat, 1975:48 [Listed ]. Parthenope serrata—Hildebrand, 1955:193.—Wass, 1955:140 [Key], 172.— Hulings, 1961:219 [Listed]_—Rouse, 1970:146. A possible means for separating the two species listed under the incertae sedis heading might be their zoogeographic distribution. Parthenope granulata is a continental species known from Cape Hatteras southward around the tip of Florida and into the eastern Gulf of Mexico as far west as Louisiana. Its excontinental distribution is Bermuda, a probable occurrence at Bahia Honda, Cuba (NMNH records see Appendix 1), and a single record from St. Thomas, Virgin Islands. It is thus possible that all specimens listed from the western Gulf of Mexico and middle and lower Caribbean southward to Brazil will prove to belong to Parthenope serrata. The latter species is known from both continental and insular records in this region. The main zone of sympatry for the two species ap- pears to be around the Florida peninsula to as far north as Cape Canaveral — on the central eastern coast. Further support for this suggestion comes from museum records on | specimens of both P. granulata and P. serrata in both the National Museum — of Natural History and the Rijksmuseum van Natuurlijke Historie in | Leiden. Dr. Lipke B. Holthuis informed me (in litt.) that in the entire © collection of “P. serrata” in the Leiden museum, six samples of P. granu- ee VOLUME 90, NUMBER 3 523 lata occur, all from stations in the Gulf of Mexico (R/V Oregon 3799, 3821, 3855, 3849, 3935, 3947). All other material in that museum, includ- ing Surinam specimens from the “Coquette” (Holthuis, 1959), and the male from Curacao taken by J. Boeke and noted by Rathbun (1919; see Appendix 1), as well as specimens collected by the University of Miami, Rosenstiel School of Marine and Atmospheric Sciences R/V John Elliott Pillsbury (stations P-330, 396, 430, 433, 444, 574) from the southwestern Caribbean Sea off Honduras, Panama, and Colombia, and R/V Oregon stations 4166 and 4179 from off the Guianas, proved to be P. serrata. Some material collected by Dr. Holthuis from Biscayne Bay, Florida, also be- longed to this species, again pointing out the zone of overlap between the two forms. Other material from the National Museum of Natural History examined by me (Appendix 2) also reflects this distribution. Acknowledgments This study could not have been completed without the gracious and generous help of many people. I wish to extend my grateful appreciation and thanks to Mr. C. Allan Child, Drs. Thomas E. Bowman, Fenner A. Chace, Jr., Raymond B. Manning, National Museum of Natural History, Washington, D.C., and to Drs. Lawrence G. Abele, Florida State Uni- versity, Tallahassee; Jacques Forest, Muséum National d'Histoire Naturelle, Paris; Bernd Hauser, Muséum d'Histoire Naturelle de Genéve, Geneva; Lipke B. Holthuis, Rijksmuseum van Natuurlijke Historie, Leiden; R. W. Ingle, British Museum (Natural History), London; Herbert W. Levi, Mu- seum of Comparative Zoology, Harvard; Austin B. Williams, National Ma- rine Fisheries Service, Systematic Laboratory, Washington, D.C.; and Torben Wolff, Universitetets Zoologiske Museum, Copenhagen. I also thank the staff of the Florida State Department of Natural Resources, Marine Research Laboratory, St. Petersburg, for making specimens of both species available for study. Photographs of the Indian River and Gulf of Mexico specimens were made by Mr. William Davenport, Harbor Branch Founda- tion, Inc. (HBF), Ft. Pierce, Florida. Those of L. crenulatus were kindly provided by Mr. G. Dajoz, Muséum d Histoire Naturelle de Geneve, through the auspices of Dr. Bernd Hauser; Dr. Jacques Forest sent the plates of L. serrata. Finally, Miss Liberta E. Scotto’s (HBF) unbounded en- thusiasm and constant good cheer relieved much of the tedium involved in this study. Her aid in literature search and with the laboratory work was invaluable. Literature Cited Aurivillius, C. W. S. 1889. Die Maskirung der oxyrrhynchen Dekapoden, durch besondere Anpassungen ihres Koérperbaues vermittelt. K. Svenska Vetenskaps Akad. Handl. 23(4):1-71, pls. 1-5. 524 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Balss, H. 1924. Westindische Decapoden. Zool. Anz. 61:177—182. Boone, L. 1927. Scientific results of the first oceanographic expedition of the “Pawnee, 1925. Crustacea from tropical east American seas. Bull. Bingham Oceanogr. Coll. 1(2):1-147, 33 text-figs. ———. 1930. Scientific results of the cruises of the yachts “Eagle” and “Ara,” 1921-1928, William K. Vanderbilt, Commanding. Crustacea: Stomatopoda and Brachyura. Bull. Vanderbilt Mar. Mus. 2:1-228, pls. 1-74. Bullis, H. R., and J. R. Thompson. 1965. Collections by the exploratory fishing vessels Oregon, Silver Bay and Pelican made during 1956-60 in the southwestern North Atlantic. Spec. Scient. Rep. U.S. Fish Wildl. Serv. (510):1—130. Chace, F. A., Jr. 1956. Crustaceos decapodos y stomatopodos del Archipiélago de Los Roques e Isla de La Orchila. In El Archipiélago de Los Roques y La Orchila. Sociedad de Ciencias Naturales La Salle, Caracas: 145-168, 4 pls. Coélho, A. P., and M. D. 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 e 39°S. Trab. Inst. Oceanogr. Univ. Recife 13:133—236, figs. 14. Desbonne, I. 1867. In: I. Desbonne and A. Schramm, Crustacés de la Guadeloupe d’aprés un manuscript du Docteur Isis Desbonne comparé avec les échantillons de crustacés de sa collection et les derniéres publications de MM. Henry de Sassure et William Stimpson. I. Partie. Brachyures [Edited, with preface, by A. Schramm]:1-60, pls. 1-8. Basse Terre. Evans, A. C. 1967. Syntypes of Decapoda described by William Stimpson and James Dana in the collections of the British Museum (Natural History). Jour. Nat. Hist. 1:399-411. Felder, D. L. 1973. An annotated key to crabs and lobsters (Decapoda, Reptantia) from coastal waters of the northwestern Gulf of Mexico. Louisiana State Unive Ruble aNos ESIW-SG=73-02-i—viis I NOSs ploy lle Flipse, H. J. 1930. Die Decapoda Brachyura der Siboga-Expedition. Part VI. Oxyrrhyncha: Parthenopidae. Siboga-Exped., Mongr. 39c?:1-96, text-figs. 1—45. —. 1931. Einige Parthenopidae aus dem naturhistorischen Museum in Leiden. Zool. Meded., Leiden 14:80-96. Godcharles, M. F., and W. C. Jaap. 1973. Fauna and flora in hydraulic clam dredge collections from Florida west and southeast coasts. Spec. Scient. Rep. Florida Dept. Nat. Res. Mar. Res. Lab. (41):1-89. Gore, R. H., and L. E. Scotto. (in press). Crabs of the family Parthenopidae (Crustacea Brachyura: Oxyrhyncha) with notes on specimens from the Indian River region of Florida. Mem. “Hourglass” Cruises, Florida Dept. Nat. Resour. Mar. Res. Lab. Gundlach, J., and J. I. Torralbas. 1889 (1900). Contribucién al estudio de los Crustaceos de Cuba. Notas compiladas y completadas por el Dr. J. I. Torralbas. An. Acad. Cienc. Méd. Fis. Nat. Habana 36:292-305. Hay, W. P., and C. A. Shore. 1918. The decapod crustaceans of Beaufort, N.C., and the surrounding region. Bull. Bur. Fish., Washington 35:369-475, pls. 25-39. Hildebrand, H. H. 1955. A study of the fauna of the pink shrimp (Penaeus duorarum Burkenroad) grounds in the Gulf of Campeche. Publs. Inst. Mar. Sci. Univ. Texas 4(1):169-232, figs. 1-2. Holthuis, L. B. 1959. The Crustacea Decapoda of Surinam (Dutch Guiana). Zool. Verh., Leiden 44:1-296, pls. 1-16, text-figs. 1-68. Hulings, N. C. 1961. The bamacle and decapod fauna from the nearshore area of Panama City, Florida. Quart. Jour. Florida Acad. Sci. 24(3):215-222. Kingsley, J. S. 1879. Notes on North American Decapoda. Proc. Boston Soc. Nat. Hist. 20:145-160. a OT VOLUME 90, NUMBER 3 52! ——. 1880. On a collection of crustacea from Virginia, North Carolina and Florida, with a revision of the genera of Crangonidae and Palaemonidae. Proc. Acad. Nat. Sci. Philadelphia 31 (for 1879):383-427, pl. 14. Leary, S. P. 1967. The crabs of Texas. Texas Parks Wildl. Dept. Coastal Fish. Bull., ser. 7, (43):1-57, plus text-figs. Martens, E. V. 1872. Ueber Cubanische Crustaceen nach den Sammlungen Dr. J. Gundlach’s. Arch. Naturgesch. 39(1):77-147, pls. 4, 5. Miers, E. J. 1879. Descriptions of new or little known species of maioid crustacea. Ann. Mag. Nat. Hist., ser. 5, 4:1-28, pls, 4, 5. . 1886. Report on the Brachyura collected by the H.M.S. Challenger during the years 1873-1876. Rep. Scient. Results H.M.S. Challenger, Zoology, 17: iio 3624 pls, 129, Milne Edwards, A. 1873-1880. Etudes sur les Xiphosures et les Crustacés de la région Mexicaine. In: Recherch. Zool. Hist. Faune Amérique Centrale Mexique, 3, By ies, ly ea —. 1880. Reports on the results of dredging, under the supervision of Alexander Agassiz, in the Gulf of Mexico, and in the Caribbean Sea, 1877, ’78, "79, by the United States Coast Survey Steamer “Blake,” Lieut.-Commander C. D. Sigsbee, U.S.N., and Commander J. R. Bartlett, U.S.N., Commanding. VIIIJ-Etudes pré- liminaires sur les Crustacés. Bull. Mus. Comp. Zool. Harvard 8(1):1-68, pls) 12) , and E. L. Bouvier. 1923. Reports on the results of dredging under the supervision of Alexander Agassiz, in the Gulf of Mexico (1877-1878) in the Caribbean Sea (1878-1879) and along the Atlantic coast of the United States (1880), by the U.S. Coast Survey Steamer “Blake” . . . XLVII. Les Por- cellanides et des Brachyures. Mem. Mus. Comp. Zool. Harv. 47(4):283-395, pls. 1-12, text-figs. 1-22. Milne Edwards, H. 1834-1840. Histoire Naturelle des Crustacés, comprenant |’anat- omie, la physiologie et la classification de ces animaux. 1:i-xxv, 1-468, and Atlas, pls. 1-42. Moreira, C. 1901. Contribucoes para o conhecimento de fauna Brazileira. Crustaceos do Brazil. Archos Mus. Nac., Rio de Janeiro 11:(v) + 1-151, pls. 1-5. Ortmann, A. FE. 1893. Die Decapoden-Krebse des Strassburger Museums. VII. Theil. Abtheilung: Brachyura (Brachyura genuina Boas) II. Unterabtheilung: Can- croidea, 2. Section: Cancrinea, 1. Gruppe: Cyclometopa. Zool. Jahrb. Abt. Syst. Okol. Geogr. Tiere, 7:411—495, pl. 17. Pequegnat, L. H. 1975. List of catalogued invertebrate species in the Texas A&M University systematic collection of marine organisms. Sea Grant Rep. TAMU- SG-76-601:iii—vi, 1-109. Rathbun, M. J. 1897. List of the decapod crustacea of Jamaica. Ann. Inst. Jamaica 1(1):1-46. —. 1898. The Brachyura of the biological expedition to the Florida Keys and the Bahamas in 1893. Bull. Labs. Nat. Hist. State Univ. Iowa 4(3):250-294, pls. 1-9. 1900. Synopses of North American Invertebrates. X. The oxyrhynchous and oxystomatous crabs of North America. American Nat. 34:503-520, 15 text-figs. 1901. The Brachyura and Macrura of Porto Rico. Bull. U.S. Fish Comm. 20 (for 1900), pt. 2:1-127, pls. 1-2, text-figs. 1-26. 1919. Stalk-eyed crustaceans of the Dutch West Indies, collected by Dr. J. Boeke 1904-1905. Rep. Fish. Aquat. Resour. Dutch W. Indies (Curacao), pt. 2:317-349, fies. 1-5. 526 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON ————. 1925. The spider crabs of America. Bull. U.S. Nat. Mus. 129:i-—xx, 1-613, pls. 1-283, text-figs. 1-153. ———. 1933. Brachyuran crabs of Porto Rico and the Virgin Islands. Scient. Sur- vey Puerto Rico 15(1):1—-121, text-figs. 1-107. 1935. Fossil Crustacea of the Atlantic and Gulf Coastal Plain. Spec. Pap. Geol. Soc. America (2):i—vii, 1-160, pls. 1-26, 2 text-figs. Righi, G. 1966. A familia Parthenopidae no Brasil (Crustacea, Brachyura). Cién. Cult., SA4o Paulo 18(2):140. Rodrigues da Costa, H. 1968. As especies de “Parthenopidae” que ocorrem nas costas do Brasil. Parte 1 (Crustacea, Brachyura). Atas Soc. Biol. Rio de Janeiro 12(3):143-147, text-figs. 1-2. —. 1969. As especies de “Parthenopidae” que ocorrem nas costas do Brasil. (Crustacea, Brachyura) Part II. Atas Soc. Biol. Rio de Janeiro 12(4):175-177, text-figs. 1-4. Rouse, W. L. 1970. Littoral crustacea from southwest Florida. Quat. Jour. Florida Acads) Seis o2(2) 2752) figs Saussure, H. de. 1858. Mémoire sur divers Crustacés nouveaux du Mexique et des Antilles. Mem. Soc. Phys. Hist. Nat. Genéve 14(2):417—496, pls. 1-6. Smith, S. I. 1886. XXI. Report on the decapod crustacea of the Albatross dredgings off the east coast of the United States during the summer and autumn of 1884. Rep. U.S. Comm. Fish. (1885), pt. 13:605-705, pls. 1-22. Springer, S., and H. R. Bullis. 1956. Collections by the Oregon in the Gulf of Mexico. 2. List of Mysidacean, Amphipod, Euphausiacean, Decapod and Stomatopod Crustaceans. Spec. Scient. Rep. U.S. Fish Wildl. Serv. 196: 1-134. Stimpson, W. 1860. Notes on North American Crustacea, in the museum of the Smithsonian Institution, No. II. Ann. Lyceum Nat. Hist. New York 7:176-246, OS, B, Be ——. 187la. Preliminary report on the Crustacea dredged in the Gulf Stream in the Straits of Florida, by L. P. de Pourtales, Assist. U.S. Coast Survey, Part 1: Brachyura. Bull. Mus. Comp. Zool. Harv. 2(2):109-160. 1871b. Notes on North American Crustacea in the Museum of the Smith- sonian Institution. No. III. Ann. Lyceum Nat. Hist. New York. 10:92-136. Tirkay, M. 1968. Dekapoden von den Margarita-Inseln (Venezuela). Senckenbergiana Biol. 49(374):249-257, plate figs. 1-8. Verrill, A. E. 1908. Decapod Crustacea of Bermuda, Part I. Brachyura and Anomura, their distribution, variations and habits. Trans. Conn. Acad. Arts Sci. 13:299- 474, pls. 9-28, text-figs. 1-67. 1922. Decapod Crustacea of Bermuda, Part II. Macrura. Trans. Conn. Acad. Arts Sci. 26:1-179, pls. 1-48, text-figs. 1-12. Wass, M. L. 1955. The decapod crustaceans of Alligator Harbor and adjacent in- shore areas of northwestern Florida. Quart. Jour. Florida Acad. Sci. 18(3): 129-176, text-figs. 1-13. White, A. 1847. List of specimens of Crustacea in the collections of the British Museum. London, i—viii + 143 pp. Williams, A. B. 1965. Marine Decapod Crustaceans of the Carolinas. Fishery Bull. U.S. Fish Wildl. Serv. 65(1):i-xi, 1-298, text-figs. 1-252. Yang, W. T. 1971. The larval and postlarval development of Parthenope serrata reared in the laboratory and the systematic position of the Parthenopinae (Crustacea, Brachyura). Biol. Bull. 140(1):166-189, figs. 1-12. Young, C. G. 1900. The stalk-eyed Crustacea of British Guiana, West Indies and Bermuda. John M. Watkins, London, 514 pp, 7 pls. ae _ VOLUME 90, NUMBER 3 527 Zeiller, W. 1974. Tropical marine invertebrates of southern Florida and the Bahama Islands. John Wiley & Sons, Inc., N.Y., i-ix + 132 pp. plus unnumbered color plates. Smithsonian Institution, Ft. Pierce Bureau, Ft. Pierce, Florida 33450. Footnotes * Scientific Contribution No. 73, from the Smithsonian Institution-Harbor Branch Foundation, Inc., Scientific Consortium, Link Port, Ft. Pierce, Florida 33450. 1 Dr. Forest noted (in litt.) “Pour les deux autres spécimens, sans étiquette, il ne sera sans doute jamais possible de prouver que ce sont des syntypes, mais a l’inverse du spécimen de Vera Cruz, il sera éqalement impossible de prouver le contraire. Le solution serait peut-etre de spécifier que, dans le cas ot la preuve serait apportee que le spécimen de Vera Cruz a été recueilli apres 1834 et que, par conséquent, il ne peut etre considéré comme syntype, il faudrait, faute d’autres syntypes certaines, le considérer comme néotype de Lambrus serratus.” *See Appendix 1. * See Appendix 1. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 528 DIOLS *q sesnyioy, AIC JfO Ant $% ‘PT (q) L668F DyNuDIs gq sesnqyioy, Ald] JO Ant 61 ¢¢ (8) L668F DIDNUDIS * J sesnyioy,, Aid JO (Ml 7p) Oks 0006 DIDS “J SeBN}.10 J, £E TA'S ‘Sn (odAyuds &) Dypjnuvis “gq Sesn}.0 J, oT 969¢¢S (pediayo uo poseg) pynjnupis *g loqieyy 9yop1eyD FO MS Ant | 9TGST (qjeuus A19 A ) dDIDLIIS * J loqieyy ay0LIeyD FO MAS Ant [| LIGSI DIDLIIS spur[sy [oqrues JO MG OG Il 6SFOS (spodouos ON) DyDjnNuDIs +g Spurs, [eqrues JjJO eT G69GG DoNUdID “dq uOIIg puUeYSsIP BIAO ST COO6F ponupis gq ay0puy JJO LS 8SPOS pDwynudis “gq loqgieyyT ayopIeyD JO MA oT LE86 pDwnuvis “g loqiefy ayopreyD Jo M PT STCST DINUDIZ “gq uonoeg Aoy ysijtoddog PT 9006F DNUdIZ “gq uolpesg Ady YON (4S T) PE LOO6F (N0ZG6F poetoquinu [eIA) DyDnNuUDIZ -g Avg Ss uewpeod Ant 2] LOT 6F ({jews Alo) gDynupIZ “gq Avg Ss ueuUIpeoc] Ant $T S616P DpnuDIS “gq Aeg suewtpeod Ant PT 6616PF DnuBIs “g [SJ 981095) 1$ Jo S Ant PT O0Z6F DIDINUDIS * J sejg ueg odey jjJO 2 IL 89ST (YqO9} UO posed) DIDjNUDID *g sefg ueg odey yO RlyD TI 9/96 DDNuDIZ “gq vloorsued oT OOSP DDINUDID “gq leat odey JJO PT S98LT DonuDIs “gq ynoyooT odey FO (OT) Owe Ol 66L8T DpnubDIs “gq setoyepyy oden FO ale YP = PUI 6VGL A}Uapl peya110H AYeIO'T suswitoods ‘ON Boleyeo WNS) ‘(QTgG ‘d ‘ojqey, :GZGT) Unqyuyey ouel Arey Aq polyHusp! vyoLLas adouay.wg Jo suauttosds Jo uoRoryQuUeploy [ xtpuoddy 529 VOLUME 90, NUMBER 3 ‘A[UO susUTToeds JYNSQ 4 ‘uees Jou ‘winesnyy Ysutig 0} osuPYyOxe U0 AT SP [y, (HIT Ul ‘sInygyOH “q “"Y) Vyas “gq ovoRiny “Avg sO.apeosol PT ‘ST Uapla'y UdVsS JON ‘ST UIsILA ‘seuoy,, 4S yay ‘sHyy Usseyusdor) DIDIMIS “d oor “qd “AqH zonseAvyy Anf 2] CESS (uasuttoeds paliq{) uses jo eoreule [ &1 ZISLI GPL8F WNSNA ([[eus A19A) gpyDjnudis “d eqny “epuoy eye Pal TAS ‘snyy DIDALIS *d IqH S.loutoAOc) ‘seuUEeyeg Ant 6] PIOTE (jews AloA) dDpDLas “J Aeg oudevosig ‘Aoy Ant 6T CLO6F DIDILIS *d [uyD yMeP ‘speoys UelOaIy (S1AO0 [) 63 ‘PT ZLOGF DIDALIS * Aeq epliopy 6 P46 6006F DIDALIS * Aeg epllopy 6¢ LIOGF DIDLIAS * Avg eplopy 66 VT OLO6F swoaM AVY JJO dT 26 TA'S ‘snyy wswaM AVY FO éT gO Sa" (peydeisojoyd avy) DyDIWas “g aM AVY JO MM GE OA COST DIDALIS “J Jeol opisul ‘JsoA\A AOY FJO ONG FOO6F owjnupis “J Jool optsur ‘soAq AVY FO Ant [ ‘anf 6T x &006F DIDAMIS *d fool opIsur YsoAq AO JO (B1A0) 6T ‘PT SO006F DIDALIS * OoIxeyy FO FIND 61 OQLO6F DIDNUDIS *g Jso\\ ADY 0} ‘JouuRyD AA Ant 6T ZOO6F 4so'T OorKeW JO FIND Ant | ELEOS DIALIS “J OOIxeyy FO FIND Cali 6668F DIDALIS “dq sesnyoy, Aiq W IW F Or (q) S668F Dypnudis “J sesnyoy, Aiq W IW F Oe (®) 9668F AJQUSp!I peye110 Ay[VIO'T suoumloods ‘ON Boleyeo WNS)D ‘ponuljuoy) ‘{[ xipueddy PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 530 ‘SOIJSIOUL Iejnsue soedeivo 10 oImjzeuLle pedijeyo uo peseq ‘satoads sity} A[qeqoig , DWLas “ LEGIT [dy g¢ ‘spueys] UIsitA “seLIOY.], “35 PT GCOSL DAIS “J OS6T WNsNYy g ‘ey ‘sesnzIOT OT 6LIGL IRATE “al SECT ounf EF ‘Bly ‘sesnj10], val GLIGL SS Se eS = NN - ‘penunuoy °g xIpueddy PROC. BIOL. SOC. WASH. 90(3), pp. 532-535 THE LARVA OF A NEW SUBTERRANEAN WATER BEETLE, HAIDEOPORUS TEXANUS (COLEOPTERA: DYTISCIDAE: HYDROPORINAE) Glenn Longley, and Paul J. Spangler The larva of the first hypogeous dytiscid, Haideoporus texanus Young and Longley (1976), from North America is described in this study. The first larva was collected by Joe Kolb in September 1974 when he was collecting aquatic organisms from a flowing artesian well for the senior author. Since that time, two additional larvae have been collected from the well. The well is located at the Aquatic Station (formerly the San Marcos Federal Fish Hatchery). A 12-inch pipe carries water from the well to a distribution chamber where a net was placed on the pipe. The net was removed and the contents were emptied into preservative approxi- mately every two days. Netting was started on a regular basis in De- cember 1973 and has continued to the present time. Twenty-nine adults of H. texanus have been collected and were described by Young and Longley (1976). In that paper there is a discussion of the ecological conditions; and, therefore, they will not be covered again in this paper. The well from which this beetle was collected was drilled in 1895 for the supply of water for the hatchery. The source of water for the well is the Edwards Aquifer, a large ground water reservoir located in Ed- wards and associated limestones along the Balcones Escarpment in Texas. Well records show that water is actually derived from a 1.5 m cavern at a depth of 59.5 m. The source for the well is the same as for San Marcos Springs, a large first order spring located approximately 670 m away from the well. This well is the type-locality for many other subterranean forms including the Texas Blind Salamander, Typhlomolge (= Eurycea) rathbuni Stejneger and several blind, depigmented invertebrates including crusta- ceans, a planarian, and a snail. Larva.—Length (including cerci), 5.5 mm; greatest width, 1.8 mm. The body is transparent with a light yellow-brown tint. Ocelli are notably ab- sent from the ocular area, resulting from an adaptation to complete sub- terranean existence (Figs. 1, 2). Head behind nasale almost quadrate. Nasale broad and reflexed (Fig. 2), elongate, anteroventral margin bearing a row of club-shaped setae; dorsal anterior margin of the nasale with long setae. Antenna cylindrical, 4 segmented; basal segment shorter than 2nd and 3rd segments; 2nd and 3rd segments about equal in length; 4th segment shortest; 3rd segment with 3 acicular setae, 1 ventrally near midlength and 2 dorsally near apex. Gular sutures barely visible (Fig. 3) on ventral surface of head. Mandible long, falciform (Fig. 2). Maxillary palpus VOLUME 90, NUMBER 3 533 3 4 Figs. 1-4. Haideoporus texanus Young and Longley, larva: 1, Head, dorsal view; 2, Head, lateral view; 3, Head, ventral view; 4, Caudal segment and cerci, dorsal view. 534 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON (Fig. 3) slender, elongate, 3 segmented; first 2 segments approximately equal in length, terminal segment shortest; 2nd segment with 2 long slender setae apicodorsally; 3rd (terminal) segment with 1 long slender seta lat- erally near apex. Stipes cylindrical and bearing 2 long setae on ventral surface, 1 basally and 1 distally. Labium setose with 5 or 6 setae on each side of the anterior margin; ligula absent; labial palpus slender, 2 segmented, terminal segment with 1 seta on inner margin at midlength. Pronotum angular at anterolateral margin, with numerous long setae at this point, also moderately long setae along lateral margin and a group of longer setae at the curved posterolateral margin. Mesonotum about half as long as pronotum, bearing numerous setae along the lateral and posterior margins. Metanotum same length as mesonotum; setation similar to that of mesonotum. Legs elongate, 5 segmented; coxa long, broad at base and tapering to apex, setose; trochanter about ’% as long as coxa, setose; femur is longest segment, 1% times longer than tibia; tibia setose and slightly shorter than tarsus; tarsus setose with 2 elongate tarsal claws. Abdomen 8 segmented, setose, with longest setae on posterior margins of abdominal segments; the last 2 segments tapering toward the tip. Cerci (Fig. 4) with 2 lateral setae arising approximately at basal 3rd, 1 lateral seta arising at apical 3rd, and each cercus with 4 long hairs on apex. The preceding description refers to a single morphotype. The speci- men was collected from the artesian well at the Aquatic Station, South- west Texas State University, San Marcos, Texas (Hays Co.) 18 Sept. 1974. The three larvae of Haideoporus texanus are fragile and because we prefer to keep the larvae intact no dissections were made for study under a com- pound microscope. Instead, the larvae were studied under a stereoscopic microscope at a magnification of 750 diameters. The larvae were compared with a paratypic larva of the Japanese Morimotoa phreatica Ueno, a blind hydroporine larva kindly provided by Dr. Ueno, and with Dr. Ueno’s (1957) description of his larva. The larvae of Haideoporus texanus are very similar to the larva of Morimotoa phreatica. The only significant differences noted are found in the antenna, the cerci, and the setae on the maxilla. The antenna of M. phreatica, according to Dr. Ueno’s illustrations (Figs. 40, 44), terminates in a small apical segment and is accompanied by a seta of similar size. However, in the larvae of M. phreatica we could detect only the small apical segment. This larva also is very fragile and could not be examined safely under a compound microscope. Because the apical segment and the seta often lie snugly side by side, we may not have been able to detect them with our stereoscopic microscope. In fact, we could see only the small apical antennal segment on the larva of M. phreatica provided by Dr. Ueno. If the seta is present on larvae of H. texanus as it VOLUME 90, NUMBER 3 535 is on most larvae belonging to the Hydroporinae, the antennae will have to be studied with a compound microscope when more larvae are obtained. The cerci of M. phreatica each bears an apparent apical segment from which 3 long setae arise at the base and one subapically. On H. texanus, each cercus (Fig. 4) is unsegmented and bears 2 lateral setae at basal 3rd, 1 lateral seta at apical 3rd and 4 apical setae, 1 of which is slightly thicker than the other 3. The last segment of the maxillary palpus of M. phreatica bears 3 subapical setae, whereas only 1 subapical seta is present on the same segment on larvae of H. texanus. The stipes of M. phreatica each bears 3 ventrobasal setae and on H. texanus, each stipes bears 2 ventral setae, 1 basal, and 1 apical. Since Haideoporus texanus was described by Young and Longley from a type-series of 29 adults, eight additional topotypic adults have been col- lected bringing the total number of adults known to 37. Adults of Haide- oporus texanus have been collected in each month of the year except August and November. One teneral adult was collected on 9 January 1974. The larva described above was collected on 18 September 1974 by Mr. Joseph Kolb. Since that time two additional larvae were collected, one on 17 August 1975 and one on 5 October 1975 by Glenn Longley. This evidence indicates that H. texanus breeds during the last 4 to 6 months of the year. Additional collecting may show whether breeding occurs throughout the year reflecting the uniform environmental conditions pres- ent in the troglobitic habitat or that a breeding season actually exists from August to January. Because all known dytiscid larvae pupate in damp places away from water it would be interesting to know where and how pupation occurs in these subterranean habitats. We surmise that the larvae pupate on the margins of the streams in the caverns which usually are only partially filled with water or in air spaces which must be trapped in large and small irregularities in the sides and ceiling of these lime- stone caverns when they may be filled with water. The three larvae will be deposited in the National Museum of Natural History, Smithsonian Insti- tution, Washington, D.C. Literature Cited Young, F. N., and G. Longley. 1976. A new subterranean aquatic beetle from Texas (Coleoptera: Dytiscidae-Hydroporinae). Ann. Ent. Soc. America 69(5):787-792. Ueno, S. 1957. Blind aquatic beetles of Japan, with some accounts of the fauna of Japanese subterranean waters. Arch. fur Hydrobiol. 53(2):250-296, figs. 1-55. Aquatic Station, Southwest Texas State University, San Marcos, Texas 78666 and Smithsonian Institution, Washington, D.C. 20560. PROC. BIOL. SOC. WASH. 90(3), pp. 536-552 THE SPHEGINA SPECIES OF EASTERN NORTH AMERICA (DIPTERA: SYRPHIDAEF) Gary A. Coovert, and F. Christian Thompson The genus Sphegina is a small group of syrphid flies that breed in sap wounds of trees. The adults are common pollinators of spring flowers in the northern forests. The nearctic species were studied or revised by Mal- loch (1922a & b), Cole (1924), and Hull (1935). The immature stages of one nearctic and three palearctic species have been described (see Laval- lee and Wallace (1973)). While working separately on revisions of the Syrphidae of Ohio (Coovert) and eastern North America (Thompson), we have discovered much new data on these flies. We present a new arrange- ment for the nearctic species, description of a new species, four new synonyms, a new key to the eastern species incorporating the previously unknown sexes of five species, and new distributional and biological data. The distributional and biological data have been condensed (only states, or provinces, and counties given); the full data have been placed in the files of the Systematic Entomology Laboratory, ARS, USDA, Washington, IDG, Stackelberg (1953) described the subgenus Asiosphegina for those species of Sphegina that lack the first abdominal sternum. While we recognize that this action leaves Sphegina proper as a paraphyletic group, we feel Asiosphegina is a useful group and accept it as valid. The nearctic species are distributed between these subgenera as follows: Asiosphegina (bi- annulata Malloch, californica Malloch, campanulata Robertson, petiolata Coquillett, and rufiventris Loew); and Sphegina (appalachiensis Coovert, armatipes Malloch, brachygaster Hull, birdwell Cole, cressoni Hull, flavi- mana Malloch, flavomaculata Malloch, infuscata Loew (= pluto Hull, new synonym), keeniana Williston, lobata Loew, lobulifera Malloch, melanderi Cole, nigrimana Cole, notata Hull, occidentalis Malloch, punctata Cole, rufa Malloch, and vittata Cole). Genus Sphegina Meigen Sphegina Meigen, 1822:193. Type-species, Milesia clunipes Fallén (West- wood, 1840:136). Small, elongate flies (4-8 mm); face strongly concave, bare; cheek linear; eyes bare, dichoptic in both sexes; antenna short, as long as face, with 3rd segment orbicular; arista bare or pubescent. Thorax: anterior mesopleuron, posterior pteropleuron, hypopleuron (including barrette), metasternum all bare; scutellum usually with marginal bristles, without subscutellar fringe; postmetacoxal bridge complete; legs normal except VOLUME 90, NUMBER 3 537 hind femur enlarged and with ventroapical spinose bristles. Wing: usually hyaline, rarely with crossveins margined with brown; alula narrow or absent, always narrower than width of anal cell; marginal cell open; apical cell closed and petiolate; anterior crossvein basal, at basal % or less of discal cell; spurious vein present or absent. Abdomen petiolate. Sphegina belongs to the subfamily Eristalinae (= Milesiinae), tribe Brachyopini (= Chrysogasterini), subtribe Spheginina (sensu Thompson, 1972:114-115). The genus is separated from all other syrphid flies by the following combination of characters: 1) postmetacoxal bridge is com- plete and broad; 2) the face is strongly concave; 3) third antennal segment is oval; and 4) the apical crossvein is oblique, forming an acute angle with the third vein. The phylogenetic relationships of Sphegina are discussed and diagrammed by Thompson (1972:114-115; 1976); the sister group to Sphegina is Neoascia Williston. In the species with an unproduced male 4th sternum or with spinose bristles on the male 3rd and 4th sterna (brachygaster Hull, flavomaculata Malloch, appalachiensis Coovert, keeniana Williston), the number of fe- males collected is greater than the number of males. All of the other species show between 1.5 and 3 times as many males collected as females, with rufiventris Loew over 6 times. Perhaps different mating behavior could ac- count for this disparity in sex ratio or else inadequate collecting techniques. Further research into the behavior of this genus of closely related flies would be rewarding, especially considering the fact that normally several different species are found flying together. Key to the subgenera of Sphegina Meigen First abdominal sternum present (fig. 16) Sphegina Meigen First abdominal sternum absent (fig. 15) Asiosphegina Stackelberg Key to the species of Sphegina (Sphegina) found in eastern North America 1. Males 2 — Females S 2. Fourth sternum laevolobate, asymmetric (figs. 2, 3) 3 — Fourth sternum not lobate, symmetric (figs. 4, 5, 8, 9, 12) 4 3. Third and 4th sterna with short, black spinose bristles apically (fig. 2); humerus pale, yellow to reddish orange; 4th sternum with lobe usually yellow lobata Loew — Third and 4th sterna with only fine pile (fig. 3); humerus black; 4th sternum with lobe usually brown to black lobulifera Malloch 4, Third and 4th sterna with short black spinose bristles apically (figs. 4, 9); sternopleuron bare and polished posterodorsally 5 Ot (dy) Co 10. ie PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Third and 4th sterna with only fine pile (figs. 5, 8, 12); sterno- pleuron completely pollinose 6 Hind tibia with apicoventral scooplike tooth; front and middle tarsi with apical 2 tarsomeres black; hind tarsus with 2nd and 3rd tarsomeres light yellow and contrasting with other darker tarsomeres keeniana Williston Hind tibia simple apically (fig. 20); front and middle tarsi with apical 2 tarsomeres yellow, but very slightly darkened; hind tarsus with 2nd and 3rd tarsomeres brown dorsally and nearly con- colorous with remaining tarsomeres appalachiensis Coovert . Seventh segment with a distinct tubercle on apicomedial margin (fig. 5); hind tibia with apicoventral scooplike tooth; 4th tergum with apical margin nearly always pallid, yellow to orange; meta- sternum and ventral 4% of postmetacoxal bridge yellow flavimana Malloch Seventh segment without a tubercle (figs. 8, 12); hind tibia simple apically or with an acute tooth; 4th tergum with apical margin black; metasternum and postmetacoxal bridge dark a Hind tibia with a small acute apicoventral tooth; vertex with pile shorter than % 3rd antennal segment width; hind femur distinctly bicolored, yellow on basal %, black on apical %; mesonotum pillinose, with two submedian shiny vittae flavomaculata Malloch Hind tibia simple apically; vertex with pile longer than % 3rd antennal segment width; hind femur not distinctly bicolored, pale on base becoming darker apically, frequently all pale; mesonotum entirely pollinose brachygaster Hull Sternopleuron bare and polished posterodorsally; hind tarsus with 2nd and 3rd tarsomeres yellow at least ventrally and contrasting with other darker tarsomeres 9 Sternopleuron completely pollinose; hind tarsus entirely dark, brown 12 Front extensively shiny, with subtriangular pollinose spots above antennae; 4th tergum entirely orange 10 Front almost entirely pollinose, with only a narrow medial shiny vitta; 4th tergum widely black apically, yellow to orange basally (humerus pale yellow to orange, contrasting with darker meso- notal disk) ‘Tel Fourth tergum flared dorsoapically (fig. 19); humerus yellow to orange, contrasting with darker mesonotal disk lobata Loew Fourth tergum not flared; humerus black, concolorous with meso- notal disk lobulifera Malloch Hind tibia with apicoventral scooplike tooth; front and middle tarsi with apical 2 tarsomeres black; hind tarsus with 2nd and VOLUME 90, NUMBER 3 539 3rd tarsomeres light yellow and contrasting with other darker tarso- meres keeniana Williston — Hind tibia simple apically (fig. 20); front and middle tarsi with apical 2 tarsomeres yellow, but very slightly darkened; hind tarsus with 2nd and 3rd tarsomeres brown dorsally and usually nearly concolorous with 3rd and 4th tarsomeres appalachiensis Coovert 12. Fifth tergum considerably broader than long, usually with apico- lateral clefts, without medial cleft (fig. 17); humerus yellow, con- trasting with darker mesonotal disk flavimana Malloch — Fifth tergum as long or longer than broad, with a single apico- medial cleft (as in fig. 18); humerus black, concolorous with mesonotal disk 13 13. Abdomen only slightly petiolate, 2nd tergum at narrowest broader than scutellum (fig. 13); vertex with pile longer than ™% 3rd an- tennal segment width; mesonotum entirely pollinose brachygaster Hull — Abdomen strongly petiolate, 2nd tergum at narrowest narrower than scutellum (as in fig. 14); vertex with pile much shorter than % 3rd antennal segment width; mesonotum at least partially shiny, shiny areas often appearing as longitudinal vittae flavomaculata Malloch Subgenus Sphegina Meigen Sphegina appalachiensis Coovert, new species Jie, DISS BAO, PAL AE BAG, a7) Small black species with or without pale abdominal markings; humerus pale in female; front extensively pollinose; mesonotum with distinct curvilinear depression extending from humerus to transverse suture; male with 4th sternum not lobate, 3rd and 4th sterna with short, black spinose bristles apically; sternopleuron bare and polished posterodorsally; hind tibia simple apically; front and middle tarsi with apical 2 tarsomeres yellow, but very slightly darkened. Male.—Length: 4.8-6.0 mm. Head: vertex black, sparse silvery gray pollinose, erect short white pilose; front black, densely silvery white pol- linose except for narrow medial triangular bare areas above lunule and below ocelli, usually meeting as a thin bare vitta, short erect white pilose laterally, with pile short and less than % 3rd antennal segment width; frontal lunule shiny black, lacking pollinose areas; antenna pale orangish yellow except first and second segments slightly darker; arista brownish yellow, darker apically, abruptly thickened on basal %s to “4, distinctly pubescent, with some of the hairs *s as long as aristal width; face dull yellow brown, black below antennae, entirely white pollinose, erect white pilose along eye margins; cheek dull yellowish brown, slightly darkened 540 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Wty Wary, “Ct ans Fig. 1. Sphegina biannulata Malloch, habitus, dorsal. at extreme eye margins, sparsely pollinose; occiput black, silvery gray pollinose, erect white pilose; eye maroon. Thorax: Entirely shiny except postalarcallus slightly brown tinged in some; dorsum short suberect pilose, silvery white pollinose anteriorly, with distinct curvilinear depression extending from humerus to trans- verse suture; pleura largely pollinose, bare and shiny on ventral % of mesopleuron, on medioventral 4% of pteropleuron and on most of sterno- pleuron posterodorsally, appressed white pilose only on posterior meso- pleuron and anterior pteropleuron; scutellum black, subappressed white VOLUME 90, NUMBER 3 541 fey e477 Figs. 2-12. Sphegina, male abdomen, ventral: 2, lobata Loew; 3, lobulifera Malloch; 4, keeniana Williston; 5, flavimana Malloch; 6, biannulata Malloch; 7, petiolata Coquillett; 8, brachygaster Hull; 9, appalachiensis Coovert; 10, rufiventris Loew; 11, campanulata Robertson; 12, flavomaculata Malloch. 542 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Figs. 13-22. Sphegina. 13-14, Scutellum and abdominal base, dorsal: 13, brachy- gaster Hull; 14, lobulifera Malloch. 15-16, Hind leg and abdominal bases, ventral; 15, biannulata Malloch; 16, brachygaster Hull. 17-18, female abdominal apex, dorsal: 17, flavimana Malloch; 18, appalachiensis Coovert. 19, lobata Loew, female abdominal apex, lateral. 20, appalachiensis Coovert, male hind leg. 21-22, aedeagus, lateral: 21, appalachiensis Coovert; 22, keeniana Williston. pilose, apical bristles present or absent. Legs: generally pale straw yellow. Following areas dark brown to black: hind femur on apical %, except extreme apices, thin ventral edge of hind tibia on basal %, hind basitarsomere, and apical 2 hind tarsomeres. Following areas brown to dark brown: subapical spot and spot at basal % on hind tibia, and 2nd and 3rd tarsomeres of hind tarsus which are slightly paler ventrally and on extreme apices. Apical 2 tarsomeres of front and middle tarsi but VOLUME 90, NUMBER 3 543 Figs. 23-28. Sphegina, male genitalia. 23-24, 9th tergum and associated structures, lateral: 23, keeniana Williston; 24, appalachiensis Coovert. 25-26, 9th tergum and associated structures, right side, dorsal: 25, keeniana Williston; 26, appalachiensis Coovert. 27-28, 9th sternum and associated structures, lateral, with a ventral view of ligula area: 27, appalachiensis Coovert; 28, keeniana Williston. slightly darkened. Pile appressed and pale. Wing: hyaline, completely microtrichose; subcostal cell yellowish brown beyond costal cell. Halter: pale yellow, brown tinged on extreme base. Squama: white with white fringe. Abdomen: Shiny black except 3rd tergum with yellow to brownish orange band on basal % which is complete, interrupted, or absent; dorsum appressed white pilose except sparsely black apicomedially on 2nd and 3rd terga; venter shiny dark brown to black except orange vitta sometimes present on 3rd sternum basally and yellow to brown apicomedially on 4th sternum; venter appressed white pilose, with numerous stubby bristle- like hairs on apicomedial areas of 2nd, 3rd, and 4th sterna. Genitalia (figs. 21, 24, 26, 27): Black, white pilose, silvery white polli- nose; cercus small, semicircular; surstyle elongate, slightly angulate apically, with a high dorsomedial carina, bare, (surstyles symmetric); 9th sternum simple, weakly sclerotized ventrad of superior lobe; ligula absent, with lig- ular area narrowly incised medially; superior lobe fused to 9th sternum, 544 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON quadrate, bare; aedeagus two segmented, with basal segment symmetric and with small apicolateral recurved spur, with distal segment asym- metric, with a small ventroapical spur on right side; aedeagal apodeme simple, small, rodshaped; ejaculatory apodeme simple, very small, rod- shaped. Female.—Length: 4.8-5.5 mm. Head: similar to that of male except 3rd antennal segment comparatively larger. Thorax: Similar to male except: humerus and postalar callus always, and area lateral to curvilinear depression usually straw yellow to yellow- ish brown; black area on hind femur confined to apical %. Abdomen: Similar to male except: pale marking present to absent on 3rd and 4th terga; 5th tergum cleft medially; pile on 2nd and 3rd terga extensively black apically; venter shiny black to mostly straw yellow, black spinose hairs absent. Types.—Holotype: ¢, WEST VIRGINIA, Pocahontas Co., Sharp Knob, 18 May 1965, 3,500 ft. (J. G. Chillcott), deposited in Canadian National Collection. Allotype: 2, same data as holotype. Paratypes: (CNC, DMNH, USNM, BM(NH)): GEORGIA: Fannin Co., Margret, 2 June 1945 (P. W. Fattig) 12. NORTH CAROLINA: Swain Co., 5 miles north Nantahalah, 8 June 1965 (J. G. Chillcott) 1¢; Mingus Cr., Cherokee-Newfound Gap, 2,000 ft., 4 June 1962 (J. G. Chillcott) 22 ?; Macon Co., Wayah Gap, 3,100 ft., 5 June 1965, on Aruncus flowers (J. G. Chillcott) 12; Wilkes Co., Doughton Gap, 2,800 ft., 6 June 1962 (J. G. Chill- cott) 12; Jackson Co., Highlands, Wilson Gap, 3,100 ft., 12 May 1957 (J. R. Vockeroth) 12; Jackson Co., Highlands, Wilson Gap, 3,100 ft., 25 May 1957 (J. R. Vockeroth) 12; Graham Co., Kilmer Memorial Forest, 1 June 1975, G. A. C. 448.53 (G. A. Coovert) 12; Graham Co., Kilmer Memorial Forest, 2 June 1975, G. A. C. 449.32 (G. A. Coovert) 12; Hay- wood Co., 27 June 1961, on Hydrangea arborescens (H. V. Weems, Jr.) 264, 722; Haywood Co., 17-19 June 1955, on Hydrangea arborescens (H. V. Weems, Jr.) 322; Haywood Co., Sunburst (O. S. Brimley) 192; Mt. Pisgah, 4-5,000 ft., 20 June 1955 & 4-8 July 1959, on Hydrangea arbore- scens (H. V. Weems, Jr.) 32 2; Dry Falls, 3 miles northwest of Highlands on Highway US 64 on Cullasaja River, 2 July 1964, on Hydrangea (H. V. Weems, Jr.) 42 2. TENNESSEE: Sevier Co., Gatlinburg-Newfound Gap, Great Smoky Mountains National Park, 3 June 1962 (J. R. Vockeroth) 22 2; Carter Co., 7 miles SW Shady Valley, 11 June 1965, on Aruncus flowers (J. G. Chillcott) 12. VIRGINIA: Floyd Co., 4 miles east Floyd, Hwy. 860, 2 June 1962, on Aruncus blossoms (J. G. Chillcott) 62 2; Patrick Co., Wool- wine, 2,500 ft., 2 June 1962, on Aruncus blossoms (J. G. Chillcott) 19; Patrick Co., Vesta, 2,800 ft., 30 May 1962, on Liriodendron blossoms (J. G. Chillcott) 12. WEST VIRGINIA: 746 6,522: same data as Holotype; Pocahontas Co., Cranberry Glades, 7-9 June 1953, on Acer spicatum (H. VOLUME 90, NUMBER 3 545 V. Weems, Jr.) 32 2; Pocahontas Co., Cranberry Glades, 1-4 June 1955, on Cornus alternifolia (H. V. Weems, Jr.) 26 ¢, 12; Pocahontas Co., Cran- berry Glades, 10 July 1967, on Hydrangea (H. V. Weems, Jr.) 32 2. Discussion This species is very similar and closely related to keeniana Williston with which it has been confused. Sphegina appalachiensis differs from keeniana Williston by the characters given in the key plus the somewhat smaller male genitalia (v. keeniana), slightly smaller black bristlelike hairs on the male abdominal sterna, and absence of a semicircular depressed area on the male fourth sternum (although this area may be weakly sclerotized and pale). As is common for species of this genus, appalachiensis flies in association with other Sphegina species, having been found with most of the other eastern species of Sphegina. Sphegina rufiventris Loew has been most commonly collected with appalachiensis which is undoubtedly due to the relative abundance of rufiventris Loew. Appalachiensis appears to be restricted to the southern portion of the Appalachian Highlands, thus the specific name. Sphegina brachygaster Hull Figs. 8, 13, 16 Sphegina brachygaster Hull, 1935:376. Type-locality: NEW YORK, Ithaca. Holotype ¢ CNC. Sphegina perplexa Hull, 1935:377. Type-locality: QUEBEC, Hull. Holo- type AMNH. ¢ New synonymy. Sphegina brimleyi Shannon, 1940:118. Type-locality: NORTH CAROLINA, Highlands, 5,000 ft. Holotype ¢ USNM. New synonymy. Distribution —Fifty-two males and 68 females examined. USA: Maine (Piscataquis), Massachusetts (Berkshire, Franklin, Hampden), New Hamp- shire (Coos, Grafton), New York (Tompkins), North Carolina (Macon, Jackson, Swain, Wilkes), Ohio (Hocking), Tennessee (Sevier), Vermont (Essex), Virginia (Giles) CANADA: Manitoba, Nova Scotia, Ontario (Simcoe), Quebec (Gatineau, Sherbrooke-Shefford). Flight period.—Earliest record 27 April (Ohio), latest record 12 July (Quebec), with most of the records in May and June. Flower records.—Daucus carota L., Lindera benzoin (L.) Blume. Variation.—Significant variation in facial color and width was noted in a long series (436 6, 322 2) collected by H. V. Weems, Jr., near Cashiers, Jackson County, North Carolina. Six males had entirely dark and broad (% of head width) faces in contrast to the normal partially yellow and narrow (6 in males, ‘40 in females, of head width) faces. While a few (5) 546 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON females also had entirely dark and broad (% of head width) faces there were some females (3) intermediate in terms of these characteristics. Comments——We have examined the holotypes of brachygaster Hull, per- plexa Hull and brimleyi Shannon and have found them to be representa- tives of the same species. We here select brachygaster Hull as senior to perplexa Hull as that name is the most appropriate descriptor for the species. Sphegina flavimana Malloch Figs. 5, 17 Sphegina flavimana Malloch, 1922a:143. Type-locality: Maryland, near Plummers Island. Holotype ¢ USNM (lost). Distribution.—Ninety-six males and 65 females examined. USA: District of Columbia, Georgia (Habersham, Rabun), Indiana (Tippecanoe), Iowa (Story), Maryland (Montgomery, Prince Georges), Massachusetts (Barn- stable, Berkshire, Hampshire, Suffolk), Michigan (Alger), New Hamp- shire (Carroll, Coos, Grafton), New Jersey (Burlington), New York (Essex, Orange, Rensselaer, Rockland, Tompkins, Suffolk, Warren), North Caro- lina (Macon, McDowell), Ohio (Adams, Ashland, Champaign, Clark, Gallia, Highland, Hocking, Lucas, Medina, Montgomery, Ross, Warren), Pennsylvania (Dauphin, Philadelphia), Tennessee (Sevier), Vermont (Ben- nington), Virginia (Fairfax, Falls Church, Floyd, Patrick), West Virginia (Ritchie, Taylor), CANADA: Manitoba, New Brunswick, Nova Scotia, On- tario (Hastings, Ottawa-Clareton, Simcoe, Thunder Bay), Quebec (Brome, Gatineau, Hull, Kamouraska). Flight period——Earliest record 2 May (Ontario), latest record 11 Sept. (North Carolina), with most of the records equally distributed between the months of May, June and July. Flower records.—Aruncus dioicus (Walt.) Fern., Ceanothus sp., Chrysan- themum sp., Cicuta maculata L., Conium maculatum L., Cryptotaenia canadensis (L.) DC., Hydrangea arborescens L., Lysimachia ciliata L. Sphegina flavomaculata Malloch Fig. 12 Sphegina flavomaculata Malloch, 1922a:141. Type-locality: VIRGINIA, Great Falls. Holotype ¢ USNM (lost). Distribution —Thirty males and 31 females examined. USA: Maryland (Montgomery), Massachusetts (Franklin, Hampden), New Hampshire (Graf- ton), New York (Orange, Tompkins), North Carolina (Graham, Macon), Ohio (Hocking, Jefferson, Pike), Pennsylvania (Philadelphia), Virginia (Fair- fax). CANADA: Quebec (Brome, Gatineau, Vercheres). VOLUME 90, NUMBER 3 547 Flight period.—Karliest record 27 March (Virginia), latest record 6 July (New Hampshire), with most of the records in May. Flower records—Caulophyllum thalictoroides (L.) Michx., Dentaria laciniata Muhl., Prunus pennsylvanica L., Rubus sp., Tiarella cordifolia L. Comments.—The characteristic apicoventral tooth on the male hind tibia, which helps to distinguish that sex, is lacking in the female. Sphegina keeniana Williston [eS 2h BR, 8) Wey 2} Sphegina keeniana Williston, 1887:113, pl. 4, fig. 11 (wing). Type-locality: PENNSYLVANIA, Philadelphia, Fairmont Park. Syntypes 36 42 (USNM, ANSP). Distribution—Ninety-three males and 118 females examined. USA: Connecticut (Fairfield, New London), Georgia (Clarke, Fannin), Indiana (Tippecanoe), Kentucky (Harlan), Maine (Hancock), Maryland (Mont- gomery, Prince Georges), Massachusetts (Berkshire, Dukes, Hampshire), New Hampshire (Coos, Grafton, Sullivan), New Jersey (Bergen, Burling- ton), New York (Bronx, Essex, Suffolk, Tompkins), North Carolina (Ma- con), Ohio (Champaign, Clark, Hamilton, Highland, Hocking, Medway, Montgomery, Pike, Ross, Summit, Vinton, Warren), Pennsylvania (Alle- gheny, Philadelphia, Susquehanna, Westmoreland), South Carolina (Pick- ens), Tennessee (Anderson), Vermont (Rutland), Virginia (Fairfax, Falls Church, Floyd, Giles, Page), West Virginia (Taylor), CANADA: Nova Scotia Ontario (Norfolk, Ottawa-Carleton, Simcoe, Wentworth, York), Quebec (Gatineau, Kamouraska). Flight period.—Earliest record 9 March (Indiana), latest record 1-15 Aug. (Vermont), with most of the records in May and June. Flower records——Aruncus dioicus (Walt.) Fern., Camassia_scilloides (Raf.) Cory, Cicuta maculata L., Conium maculatum L., Cryptotaenia canadensis (L.) DC., Daucus carota L., Galax aphylla L., Geranium sp., Heracleum maximum Bartr., Hydrangea arborescens L., Rubus sp., Sanic- ula marilandica L. Comments.—This species can only be confused with appalachiensis from which it can be separated by the apicoventral scooplike tooth on the hind tibia and the structure of the male genitalia. This tooth, unlike that of flavomaculata Malloch, is present in both sexes, but is more prominent in the male. The male genitalia of keeniana differ from those of appala- chiensis as follows: 1) surstyle is bare, with a smaller and lower dorso- medial carina, and with apex not as strongly angulate; 2) superior lobe is triangular, not quadrate; 3) ligula is broad, and convex apically, not absent: 4) aedeagal apodeme is enlarged anteriorly; 5) basal segment of aedeagus 548 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON has a dorsomedial projection as well as a larger apicolateral spur; and 6) distal segment of aedeagus lacks an apicolateral spur. Sphegina lobata Loew Figs. 2, 19 Sphegina lobata Loew, 1863:12. Type-locality: “Mittelstaaten.” Syntypes VS 2 MEZ. Sphegina monticola Malloch, 1922a:142. Type-locality: NEW HAMP- SHIRE, Mount Washington. Holotype 2 USNM. New synonymy. Distribution.—Sixty-eight males and 28 females examined. USA: Maine (Penobscot, Piscataquis), Michigan (Alger), New Hampshire (Coos, Grat- ton), New York (Essex), North Carolina (Avery, Graham, Haywood, Ma- con, Swain), Ohio (Pike), Tennessee (Sevier), Vermont (Bennington, Essex), Virginia (Smyth). CANADA: Ontario (Simcoe, Wellington), Quebec (Brome, Gatineau, Hull, Temiscamingue). Flight period Earliest record 16 May (Quebec), latest record 6 Aug. (Quebec), with most of the records in June and July. Flower records.—Aralia nudicaulis L., Chrysanthemum sp., Clintonia borealis (Ait.) Faf., Cryptotaenia canadensis (L.) DC., Daucus carota L.., Hydrangea arborescens L., Osmorhiza sp., Rubus sp., Viburnum recognitum Fern. Comments.—The holotype of monticola Malloch has been examined and found to be a female of lobata Loew, a form unknown to Malloch. Sphegina lobulifera Malloch Figs. 3, 14 Sphegina lobulifera Malloch, 1922b:269. Type-locality: MARYLAND, Plummers Island. Holotype ¢ USNM (lost). Distribution—Fifteen males and 5 females examined. USA: Connecticut (Fairfield, Litchfield), Maryland (Montgomery, Prince Georges), Massa- chusetts (Bristol), Mississippi (Lafayette), New Hampshire (Grafton, Coos), North Carolina (Avery, Macon), Ohio (Jefferson), Gaspe, Vermont (Essex), Virginia (Fairfax, Giles). CANADA: Quebec (Gaspe, Kamouraska, Que- bec). Flight period.—Earliest record 1-15 Apr. (Mississippi), latest record 10 July (Quebec), with most of the records in June. Flower records——Aruncus dioicus (Walt.) Fern., Caulophyllum thalic- troides (L.) Michx., Heracleum maximum Bartr., Rubus sp. Comments.—As evidenced by the numbers of specimens collected, this is the least common eastern species. VOLUME 90, NUMBER 3 549 Key to the species of Sphegina (Asiosphegina) found in Eastern North America 1. Mesonotum entirely shiny on disk; front shiny on dorsal %; arista pubescent, with hairs longer than aristal width; male 4th sternum with weakly sclerotized pallid lobe apicolaterally and discontinuous with sternum (figs, 6, 7) 2D, — Mesonotum with 2 broad medial pollinose vittae; front pollinose at least laterally on dorsal half; arista nearly bare, hairs much shorter than aristal width; male 4th sternum produced triangularly on left side, with lobe continuous and concolorous (figs. 10, 11) 3 2. Scutellum with a pair of long apical bristles; front and middle tarsi with apical 2 tarsomeres black; male 4th sternum with large apico- lateral lobe on left side (fig. 7) petiolata Coquillett — Scutellum without bristles; front and middle tarsi with apical 2 tarsomeres yellow to slightly brownish; male 4th sternum with a narrow submedial shelflike lobe (fig. 6) biannulata Malloch 3. Third antennal segment partially to entirely orange; wing com- pletely microtrichose; male 4th sternum without black spinose bristles, asymmetric, produced apicolaterally on left side (fig. 11) campanulata Robertson — Third antennal segment dark brown to black; wing partially bare basally, 2nd basal cell bare on anterobasal “5; male 4th sternum usually with short black spinose bristles medially, not strongly asymmetric (fig. 10) rufiventris Loew Subgenus Asiosphegina Stackelberg Sphegina, subgenus Asiosphegina Stackelberg, 1974:446 (1953:376). Type- species, Sphegina sibirica Stackelberg (Stackelberg, 1974:446) NB: The name Asiosphegina is available from 1974 when Stackelberg designated a type-species, not 1953 when he described the subgenus. Sphegina biannulata Malloch Hie sagem Gvelli Sphegina biannulata Malloch, 1922a:143. Type-locality: VIRGINIA, near Plummers Island. Holotype 2 USNM (lost). Distribution—Twenty-seven males and 14 females examined. USA: Georgia (Rabun), North Carolina (Graham, Jackson, Macon), Pennsylvania (Philadelphia), Tennessee (Sevier), Virginia (Fairfax). Flight period.—Earliest record 18 May (Tennessee), latest record 10 Aug. (North Carolina), with most of the records in May and June. Flower records.—-Galax aphylla L. 550 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Sphegina campanulata Robertson Fig. 11 Sphegina campanulata Robertson, 1901:284. Type-locality: ILLINOIS, Carlinville. Syntypes 2¢ INHS. Distribution Seventy-four males and 31 females examined. USA: Con- necticut (Hartford), Illinois (Macoupin), Indiana (Tippecanoe), Iowa (Story), Maine (Franklin, Piscataquis), Maryland (Montgomery, Prince Georges), Missouri (Jackson), New Hampshire (Cheshire, Sullivan), New York (Greene), North Carolina (Graham, Jackson, Macon, Swain), Ohio (Athens, Clark, Greene, Hamilton, Hocking, Montgomery, Pike, Scioto, Warren), Pennsylvania (Allegheny, Philadelphia), Tennessee (Sevier), Ver- mont (Bennington), Virginia (Fairfax, Floyd, Giles), West Virginia (Ritchie, Taylor), CANADA: Ontario (Essex, Ottawa-Carleton, Simcoe), Quebec (Gatineau). Flight period.—Earliest record 3 May (Indiana), latest record 1 Aug. (Virginia), with most of the records from mid-May to mid-July. Flower records.—Aruncus dioicus (Walt.) Fern., Cryptotaenia canadensis (L.) DC., Hydrangea arborescens L., Hydrophyllum sp., Osmorhiza sp., Rubus sp., Sambucus sp., Viburnum recognitum Fern. Sphegina petiolata Coquillett lanier, Sphegina petiolata Coquillett, 1910:125. Type-locality: NEW HAMP- SHIRE, White Mountains. Holotype ¢ USNM. Distribution.—Forty-six males and 23 females examined. USA: Maine (Hancock), Maryland (Prince Georges), Massachusetts (Hampshire, Wor- cester), New Hampshire (Coos, Grafton), New Jersey (Bergen), New York (Essex, Tompkins), North Carolina (Macon), Ohio (Ashland, Champaign), West Virginia (Pocahontas), Wisconsin (Douglas). CANADA: Manitoba, New Brunswick, Ontario (Norfolk, Oxford, Simcoe, Sudbury), Quebec (Gati- neau, Joliette, Kamouraska, Papineau, Terrebonne). Flight period Earliest record 18 May (W. Virginia), latest record 27 August (Maine), with most of the records from June to mid-August. Flower records.—Blephilia sp., Chrysanthemum leucanthemum L., Cryp- totaenia canadensis (L.) DC., Sanicula marilandica L., Veratrum sp. Sphegina rufiventris Loew Fig. 10 Sphegina rufiventris Loew, 1863:13. Type-locality: NEW YORK. Syntypes | MCZ. Distribution —Five hundred eight males and 77 females examined. USA: | Connecticut (Fairfield), District of Columbia, Georgia (Rabun), Illinois | | L VOLUME 90, NUMBER 3 551 (Carroll), Kentucky (Harlan), Maine (Franklin, Oxford, Penobscot, Piscata- quis, Washington), Maryland (Frederick, Montgomery), Massachusetts (Mid- dlesex, Worcester), New Hampshire (Carroll, Cheshire, Coos, Grafton), New Jersey (Burlington, Passaic, Sussex), New York (Essex, Orange, Tompkins), North Carolina (Avery, Graham, Haywood, Jackson, Macon, Swain, Wilkes, Yancey), Ohio (Champaign, Clark, Franklin, Gallia, Highland, Hocking, Montgomery, Pike), Pennsylvania (Allegheny, Carbon/Lehigh, Lycoming, Philadelphia, Westmoreland), Tennessee (Anderson, Campbell, Sevier), Ver- mont (Essex, Orleans, Rutland), Virginia (Arlington, Augusta, Buchanan, Fairfax, Falls Church, Giles, Page, Patrick, Rockbridge, Smyth, Sussex), West Virginia (Monongalia, Pendleton, Ritchie, Taylor) CANADA: New Brunswick, Newfoundland, Ontario (Northumberland, Ottawa-Carleton, Simcoe), Quebec (Gaspe-este & ouest, Gatineau, Joliette, Kamouraska, Lac-St. Jean-Ouest, Megantic, Saguenay, Sherbrooke-Shefford, Vercheres). Flight period.—Earliest record 27 April (Maryland), latest record 13 Aug. (Quebec), with most of the records in June and July. Flower records.—Aralia nudicaulis L., Aruncus dioicus (Walt.) Fern., Castanea pumila (L.) Mill., Ceanothus sp., Clintonia borealis (Ait.) Raf., Conium maculatum L., Cornus canadensis L., C. stolonifera Michx., Crypto- taenia canadensis (L.) DC., Daucus carota L., Heracleum sp., Hydrangea arborescens L., Prunus virginiana L., Rubus sp., Saxifraga micranthidifolia (Haw.) Britt., Spiraea tomentosa L., Viburnum recognitum Fern., Viburnum sp., Washingtonia sp. Comments.—This is by far our most common species, being found more commonly in the summer months. The number of males collected out- number the females by over 6 times, being a much higher ratio than any other Eastern Sphegina. Acknowledgments We especially thank the Ohio Biological Survey for their support of the work on Ohio Syrphidae which has yielded much of the new information in this paper; we also thank Charles A. Triplehorn of Ohio State Univer- sity, Columbus, J. R. Vockeroth of the Biosystematic Research Institute, Canada Department of Agriculture, Ottawa, and Howard V. Weems, Sr.., Florida State Collection of Arthropods (Syrphids from his personal col- lection), Florida Department of Agriculture and Consumer Services, Gaines- ville, for their permission to study the material in their care. Figures 1-20 were drawn by Holly K. Coovert and figures 21-28 were drawn by the junior author; the manuscript was typed by Betty J. Thompson. Literature Cited Cole, F. R. 1924. Notes on Diptera of the syrphid genus Sphegina. Entomol. News 35:39-44, 1 pl. 502 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Coquillett, D. W. 1910. New genera and species of North American Diptera. Proc. Entomol. Soc. Washington 12:124-131. Hull, F. M. 1935. Descriptions of new species of the genus Sphegina with a key to those known from North America (Syrphidae: Diptera). Trans. Amer. Entomol. Soc. 61:373-382, 1 pl. Lavallee, A. G., and J. B. Wallace. 1974. Immature stages of Milesiinae (Syrphidae). Il: Sphegina keeniana and Chrysogaster nitida. J. Georgia Entomol. Soc. 9:8-15, 24 figs. Loew, H. 1963. Diptera Americae septentrionalis indigena. Centuria tertia. Berlin. Entomol. Ztschr. 7:1—55. Malloch, J. R. 1922a. Seven new species of the syrphid genus Sphegina Meigen (Diptera). Proc. Biol. Soc. Washington 35:141-144. 1922b. Keys to the syrphid genus Sphegina Meigen (Dip.) Entomol. News 33: 266-270. Meigen, J. W. 1822. Systematische Beschreibung der bekannten europaischen zwei- fliigeligen Insekten. Vol. 3, x + 416 pp., pls. 22-32. Hamm. Robertson, C. 1901. Some new Diptera. Canad. Entomol. 33:284-286. Shannon, R. C. 1940. Highland Syrphidae (Diptera) of North Carolina. Proc. En- tomol. Soc. Washington 42:117—-120. Stackelberg, A. A. 1953. [A short survey of the Palaearctic species of the genus Sphegina Mg. (Diptera, Syrphidae).] Trudy Zool. Inst. Akad. nauk SSSR 13: 373-386, 34 figs. [in Russian, with German Summary]. ——. 1974. [New species of hover flies (Diptera, Syrphidae) of Siberia and Mongolian People’s Republic.] Entomol. Obozr. 53:443-446 [in Russian]. Thompson, F. C. 1972. A contribution to a generic revision of the neotropical Milesinae (Diptera: Syrphidae). Arq. Zool. 23:73-215, 74 figs., 11 maps. ———. 1976. Austroascia segersi, a new genus and species from Chile (Diptera: Syrphidae). Papeis Avulsos Zool. (In Press.) Westwood, J. O. 1840. Order XIII. Diptera Aristotle (Antliata Fabricius. Halterip- tera Clairv.). Pp. 125-128 in his An introduction to the modern classification of insects. Synopsis of the genera of British insects, 158 pp. London. Williston, S. W. 1887. Synopsis of the North American Syrphidae. Bull. U.S. Natn. Mus. 31:i—xxx, 1-335, 12 pls. (1886). The Dayton Museum of Natural History, Dayton, Ohio and Systematic Entomology Laboratory, ARS, USDA, Washington, D.C. Mailing addresses—(GAC) 2629 Ridge Avenue, Dayton, Ohio 45414; (FCT) c/o U.S. National Museum, NHB-168, Washington, D.C. PROC. BIOL. SOC. WASH. 90(3), pp. 553-565 KEY TO THE NEOTROPICAL GENERA OF PARYDRINAE WITH A REVISION OF THE GENUS ELELEIDES CRESSON (DIPTERA: EPHYDRIDAE) Wayne N. Mathis Abstract—Mathis, W. N., Department of Entomology, National Mu- seum of Natural History, NHB 169, Smithsonian Institution, Washington, D.C. 20560.—Eleleides Cresson is revised to include one new species, E. liroceras (type-locality, Hurlingham, Buenos Aires, Argentina). Characters of the head, thorax, and male and female postabdomens are illustrated. The genus is placed in the tribe Hyadinini, subfamily Parydrinae. Other Neotropical genera of Parydrinae are keyed and their relationships dis- cussed. Shore flies (ephydrids) of the Neotropics are very poorly known. Many undescribed taxa presently await description and revisionary study and numerous others undoubtedly remain to be discovered. Consequently, the classification of this fauna is in a preliminary state, subject to considerable change and revision with the addition of new taxa. The most recent cata- logue of Neotropical Diptera (Wirth, 1968) lists three tribes, 11 genera, and 41 species for the subfamily Parydrinae. With few exceptions, most of these taxa were described in isolated descriptions, lacking keys or illustra- tions. Exceptions are the studies of Cresson (1931) on the fauna of south- ern Chile and Patagonia; Cresson (1934) on the genus Beckeriella; Cresson (1938) on the genera Ochthera and Stenochthera; Wirth (1958) on the genus Gastrops; and Wirth (1970) on the genus Physemops. The purpose of the present paper is to provide a key to Neotropical genera of Parydrinae, to make some preliminary observations regarding the phylogeny and classification of the Central and South American fauna of this subfamily, and to revise the genus Eleleides Cresson. Genera included in the key are those listed in the aforementioned cata- logue with the addition of Pelina and Diedrops. A species of Pelina is now known to occur in the Neotropics (Clausen, 1973) and Diedrops was described from Mexican and Peruvian specimens since the publication of the catalogue (Mathis and Wirth, 1976). Key to the genera of Neotropical Parydrinae (Ephydridae) 1. Eye pilose; facial height at middle subequal to that of clypeus; antenna small, inserted in widely separated cavities; arista rudi- mentary, length when visible approximately ‘2 width of 3rd an- bo Ol PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON tennal segment; head and mesonotum mostly bare, lacking any prominent bristles; generally cinereous and dorsoventrally flat- tened Lipochaeta Coquillett Eye bare; face much higher than clypeus; antennal bases approxi- mate, arista normally developed; head and mesonotum usually with some distinguishable bristles; shape and coloration not as above 2 Face vertically carinate; costa extending to vein R 3 + 4; vein R 2 + 3 short, merging with costa before level of posterior crossvein; arista pectinate; aristal branches 7 to 11, longer ones equal to length of 3rd antennal segment Brachydeutera Loew Face flat to convex, not vertically carinate; costa extending to vein M 1 + 2; vein R 2 + 3 longer, merging with costa at or beyond level of posterior crossvein; arista variable, bare or pectinate, if pectinate with fewer branches or longer branches not equal in length to 3rd antennal segment 3 Oral opening large, gaping; clypeus prominent as a transverse band Parydra Stenhammer Oral opening small to moderately large; clypeus variously exposed, usually tonguelike, not as transverse band 4 Arista pectinate; mid tibia with a prominent ventroapical bristle 5 Arista bare to macropubescent; mid tibia lacking a promient ventro- apical bristle 9 Aristal branches 3 to 4; wing hyaline; abdominal terga 2-4 subequal in width, pollinose but more or less smooth; halter capitellum pale 6 Aristal branches 7 or more; wing usually infuscated often with dis- tinct maculation pattern; abdominal terga 3 and 4 enlarged, wider than 2, rugose or pitted; halter capitellum black 8 Fore leg normal, with femur and tibia similar to those of mid and hind legs; chaetotaxy of head and mesonotum well developed; pos- terior crossvein perpendicular to vein M 1 + 2 Eleleides Cresson Fore leg raptorial, with femur greatly enlarged and spinose below, tibial apex produced ventrally into a long, stout, extending spur; chaetotaxy of head and mesonotum reduced, mostly lacking; pos- terior crossvein merging with vein M 1 + 2 at acute angle ih Eye much higher than wide; gena very narrow, usually less than % eye height; face relatively narrow, especially below antennal fovea, not vertically carinate or tuberculate Stenochthera Hendel Eye only slightly higher than wide; gena wide, approximately % eye height; face relatively wide, vertically carinate or tuberculate Ochthtera Latreille Face flat or concave Beckeriella Williston Ul Ol Ot VOLUME 90, NUMBER 3 — Face with gently rounded, broad swelling on dorsal half Gastrops Williston 9. Face broadly arched, shieldlike, sparsely setulose; clypeus con- cealed 10 — Face flat to convex but not shieldlike, mostly bare; clypeus variable, if exposed, tonguelike 11 10. Vein R 2 + 3 very long, ending in costa at about same level as R 4 + 5; arista shorter, length much less than width of frons just above antennae; halter discolored, dark but not distinctly black; larger species, usually more than3 mm _—_ Diedrops Mathis and Wirth — Vein R 2 + 3 shorter, ending in costa before R 4 + 5; arista longer, length subequal to width of frons just above antenna; halter capi- tellum black; small species, less than 3 mm Physemops Cresson 11. Vein R 2 + 3 longer, costal vein ratio 1:0.5 or less 12 — Vein R 2 + 3 shorter, costal vein ratio 1:0.8 or more i 12. Gena large, height greater than % eye height; only inner vertical bristle present; mostly brown, pollinose species. _—_ Pelionoides Cresson — Gena smaller, height much less than % eye height; 1 pair of fronto- orbital bristles and both inner and outer vertical bristles present; mostly black, subshiny to shiny species Pelina Haliday 13. Fourth abdominal tergum longer than combined length of 2nd and 3rd terga; only inner vertical bristle evident Lytogaster Becker — Fourth abdominal tergum shorter, not longer than combined length of 2nd and 3rd terga; vertical bristles variable, often outer present Hyadina Haliday While constructing the key to Neotropical genera of Parydrinae, sey- eral problems became apparent that need further commentary. My com- ments must be considered preliminary in the sense that the problems addressed are not resolved beyond the suggestion of possibilities in most instances. My reasons for commenting now are to bring the problems to the forefront, to provide perspective, to point out the vast lacuna in our understanding of the Neotropical shore fly fauna, and to encourage further collection and study of Central and South American ephydrids. Physemops and Diedrops are included in the key because recent studies (Wirth, 1968 and Mathis and Wirth, 1976, respectively) have assigned these genera to the subfamily Parydrinae. However, members of these genera plus those of Dagus Cresson (Neotropical), Psilephydra Hendel (Oriental), and Apulvillus Malloch (Polynesian) share many similarities (outlined below) and will probably form the basis of a taxon inter- mediate between Ephydrinae and Parydrinae or incorporated within the enlarged concept of either subfamily. Wirth (1970) correctly pointed out that Physemops and Psilephydra are similar to Apulvillus, a genus of the 5906 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON tribe Scatellini (Ephydrinae). I have only added Dagus and Diedrops to the genera included in the group (hereafter denoted as Physemops-group). The face of members of these genera is projecting or shieldlike, concealing the clypeus, is arched, sparsely setulose, and is invested with silvery- white pubescence. Also, members of these genera have some, although often reduced, mesonotal (dorsocentral and acrostichal) and cephalic bristles, have a characteristically shortened frons, and aside from specimens of Physemops, have dull to subshiny-brown coloration dorsally and are contrastingly gray below with a rather sharp demarcation between the coloration change. The dorsal surface is usually dull, often pollinose, but is uniformly smooth. Specimens of Physemops differ by being entirely dark brown to black and shinier. This brief outline of characters contrasts with those of most genera of Parydrinae (s. str.) which are subshiny to shiny and have the cuticular surface, notably on the mesonotum, granulose to microrugose and for the most part lacking prominent bristles. A direct relationship between the Physemops-group and Parydrinae (s. str.) is not established. However, the genus Brachydeutera is possibly intermediate. Brachydeutera specimens have coloration and cuticular sur- face similarities with members of the Physemops-group but also resemble those of the tribe Parydrini (Parydrinae), i.e. clypeus exposed, transverse and oral opening large, gaping. A second possible connection between the Physemops-group and Parydrinae is through Eleleides, the genus revised below. Members of Eleleides, like Brachydeutera are similar to those of the Physemops-group in coloration and surface texture; they also possess several well-developed dorsocentral and fronto-orbital bristles. Like other parydrine genera, especially Ochthera and Stenochthera, the clypeus of Eleleides specimens is exposed, appearing tonguelike through the ventral facial emargination. In addition, the overall appearance of Eleleides speci- mens is similar to that of Pelinoides specimens. From the above, it is obvious that the relationships of the Physemops-group with other ephydrid taxa are poorly known. An evaluation of pertinent characters, to determine their relative derived versus primitive states will be necessary before the sequence of lineages can be better clarified. Ochthera and Stenochthera members are closely related as demonstrated by many synapomorphies (second half of key couplet 6). However, recog- nition of each as a distinct genus is questionable. Stenochthera was orig- inally proposed by Hendel (1930), who used characters that are now known to vary considerably within members of Ochthera alone. Although Stenochthera specimens can generally be recognized as a distinct taxon in the Neotropics, there are few characters that are reliable for recognizing the group on a worldwide basis. However, the surstyli of the few Stenoch- thera species I examined were somewhat unique, having a median con- striction, superficially dividing each surstylus into 2 parts. —~ VOLUME 90, NUMBER 3 507 My comments questioning the generic distinctiveness of Ochthera and Stenochthera also apply to Beckeriella and Gastrops. The only consistent character to distinguish between the latter 2 genera is the concave to slightly convex face of Beckeriella specimens as opposed to the upper facial tumescence in members of Gastrops. Cresson (1934:201) while re- viewing the species of Beckeriella, also noted the close similarity between members of these genera, stating that “The structural similarity between the species of this genus and those of Gastrops is very marked, in fact, were it not for the flat to concave or slightly convex face of Beckeriella as opposed to the characteristically strongly convex to gibbose face of Gastrops, the two groups of species could almost be considered congeneric. The general structure of the head, thorax and abdomen is so similar in the two genera that their close relationship is evident.” Fifteen years later, Cresson (1949) erected the tribe Gastropsini for these genera based on the distinc- tive conformation of their heads and pectinate arista. This tribe has not since been generally recognized (Wirth, 1968). Regardless of generic status however, these taxa are a monophyletic lineage as established by the synapo- morphies cited in the key (second half of couplet 5). Further classification of the relationships suggested above will depend on in depth studies of the genera in question. Undoubtedly there are many characters of the male and female postabdomen that will add consider- ably to a better understanding of the subfamilial and generic relation- ships. Eleleides Cresson Eleleides Cresson, 1948:20. Type-species: Eleleides chloris Cresson, by original designation and monotypy. Wirth, 1968:141-142 (review). Diagnosis—Members of Eleleides may be distinguished from those of similar genera of the subfamily Parydrinae by the following combination of characters: apical, spinelike seta of second antennal segment small, inconspicuous; arista with 3 to 5 branches above; eye bare; oral opening small; face with ventral margin deeply emarginate at middle; clypeus ex- posed, protruding through facial emargination; face much narrower than frons, mostly flat, at least lower portion densely pollinose; 1 pair of well- developed, crucinate facial bristles; 1 pair of larger fronto-orbital bristles with lateral posteroblique orientation; dorsocentral bristles well developed. 2 pairs (1 + 1); fore legs normally developed, similar to mid and hind legs; fore femur with anteroventral series of minute, closely spaced, short. flattened denticles; mid tibia lacking dorsal erect extensor bristles; wing hyaline; posterior crossvein perpendicular to vein M, 2; most of abdomen shiny, with metallic green to dark bluish-green coloration. Description—Small ephydrids, length approximately 2 mm; generally 558 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON dark colored above, pollinose to shiny; pleural areas contrastingly lighter in coloration, distinctly pollinose. Head: Slightly wider than high; frons wider than long, trapezoidal in shape becoming narrower anteriorly; mesofrons not obviously distinct from parafrons in color or texture; ocelli arranged in isoceles triangle, distance be- tween posterior pair longer than between either posterior ocellus and median ocellus; chaetotaxy of frons as follows: proclinate fronto-orbitals 2 pairs, posterior pair much smaller; lateroclinate fronto-orbital large, inserted me- dially to proclinate fronto-orbitals, close to posterior fronto-orbital; both inner and outer vertical bristles well developed, distance between them less than between inner vertical and posterior fronto-orbital, the latter dis- tance subequal to distance between posterior ocelli; 1 pair of large, pro- clinate, parallel ocellar bristles; dorsalmost pair of postocular bristles more prominent, stronger, several smaller bristles along posterior margin of eye. Second antennal segment with a short bristle above and several less evident setae on ventral and median surfaces; third antennal segment as wide as long; arista slightly enlarged basally, macropubescent, stylelike apically, with 3-4 branches above. Face flat in profile, higher than width between eyes, becoming considerably broader ventrally; uniformly textured, densely pollinose, and colored; deeply emarginate along ventral edge at middle, exposing clypeus and maxillary palps; facial bristles large, crucinate, 1 pair at level of ventral margin of eye, otherwise face mostly bare. Para- facials gradually becoming wider ventrally. Eye bare, almost as wide as high, suboval, oriented at slight oblique angle to epistoma. Gena wide to narrow, bearing 1 large bristle and several smaller ones toward postero- ventral edge of head; maxillary palp spatulate; oral opening small, pre- mentum narrow, shiny. Thorax: Dark colored above, pollinose to shiny, becoming shinier pos- teriorly; pleural areas distinctly contrasting with mesonotum, lighter, more pollinose. Thoracic chaetotaxy as follows: acrostichal bristles generally in- conspicuous except for 1 pair of widely separated prescutellars, occasionally with | larger pair at sutural level; 2 pairs of dorsocentral bristles (1 + 1): 1 pair of presutural bristles; 1 or 2 pairs of supra-alar bristles; 1 pair of intra- alar bristles; 1 pair of humeral bristles; 2 pairs of notopleural bristles, both inserted at same distance from ventral notopleural margin; 2 pairs of larger lateral scutellars, apical pair slightly larger, with 1 pair of much smaller bristles between larger scutellars; 2 pairs of larger mesopleural bristles along posterior margin; 1 pair of sternopleural bristles; otherwise pleural areas mostly bare. Legs more or less subequal to each other, normal, femora not greatly enlarged; mid tibia with prominent ventroapical bristle. Wing hyaline; costal vein extending to vein M,.»2. Halter pale. Abdomen: Shiny dorsally with metallic reflections; terga 2-4 sub- VOLUME 90, NUMBER 3 599 equal in length, fifth slightly shorter. Female with 6 visible terga; segments 1-7 complete but with distinctly larger membraneous gap between seg- ments 6 and 7 than between first 6 segments. Ventral receptacle with oper- culum much higher than wide, extending process simple. Male genitalia symmetrical, cercus prominent, long; posterior surstylus large, attached but not fused indistinguishably with ventral margin of epandrium; anterior surstylus prominent, setulose; aedeagus wide basally, sometimes terminat- ing with long crescent-shaped, narrow process. Geographic distribution —Disjunct; southeastern Australia and north- eastern Argentina. I suspect that with more thorough collecting, the genus will be found to be more widespread than is presently indicated and that ad- ditional species will be discovered. Moreover, I suggest that western South America and possibly temperate Africa will be productive in this regard. Based on the known disjunct distribution, however, the possibility that one or the other species was introduced must also be considered. Discussion—Members of the genus Eleleides have close affinities with genera of the tribe Hyadinini sensu Wirth and Stone (1956) of the sub- family Parydrinae and are being placed in this tribe, despite similarities Eleleides has with genera of Typopsilopini (Notiphilinae), the tribe in which Cresson (1948) originally described the genus. The relationship of Eleleides with Hyadinini is evidenced by the following character states, exhibited in Eleleides specimens and also used to recognize the tribe: eye bare; antenna normally developed; arista present, well developed, with 3-5 branches above; 1 larger pair of fronto-orbital bristles with a later- oblique orientation; oral opening small to moderately large; and clypeus exposed through facial emargination, but not prominent as a large trans- verse band. Within the tribe Hyadinini, Eleleides appears to be allied with Ochthera and Stenochthera and it is probably the sister-group of the latter genera and perhaps to the remaining genera of Hyadinini. The relationship be- tween members of Ochthera + Stenochthera and those of Eleleides is based on the joint possession of sparsely branched, pectinate arista; the occurrence of a rather large, pendulous posterior surstylus that is not fused to the ventral margin of the epandrium; the row of cuticular, peglike denti- cles along the venter of the fore femur, and the general features of the head, especially the deeply emarginate ventral edge of the face through which the clypeus protrudes. Specimens of Eleleides may be readily dis- tinguished from either Ochthera or Stenochthera by the unusually well- developed chaetotaxy of the head and thorax, particularly the large fronto-orbital and prominent dorsocentral bristles and by the normally de- veloped fore leg. The strongly developed chactotaxy is unique among most Parydrinae genera and distinguishes specimens of Eleleides from all others. 560 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Eleleides liroceras, new species Figs. 1-10 Diagnosis —Specimens of E. liroceras are distinguished by the following combination of characters: eye-to-cheek ratio more than 1:0.40; third an- tennal segment pale, yellowish orange to orange; frons subshiny, with bronzish-brown, pollinose vestiture except for cinereous anterior margin; face unicolorous, gray to silvery gray; maxillary palp pale, yellowish orange; anterior portion of mesonotum dull, concolorous with frons, becoming sub- shiny to shiny, darker, more greenish blue posteriorly; bristles in general more strongly developed; no larger pairs of acrostichal bristles; anterior supra- alar bristle lacking; tibiae pale, concolorous with basal tarsomeres, con- trasting distinctly with darker femora; abdominal terga with lateral cinere- ous wedges, these contrasting with subshiny to shiny areas. Description Length 1.84 to 2.48 mm (averaging 2.16 mm). Head (Figs. 1 and 2): Head width-to-height ratio averaging 1:0.78; frons width-to-length ratio averaging 1:0.60; frons mostly subshiny black with dense bronzish-brown pollinose vestiture that becomes contrastingly gray along anterior margin. Fronto-orbital plates and mesofrons slightly raised in relief and shinier. First and second antennal segments dark brown, dull; third segment as wide as long, entirely pale, orange, macropubescent along rounded apical edge. Face and gena concolorous, entirely gray to silvery gray, face height-to-width ratio averaging 1:0.60; clypeus black, not as gray pollinose as face; maxillary palp entirely pale yellow. Eye height-to-width ratio averaging 1:0.94; eye-to-cheek ratio averaging 1: 0.42; ventral portion of gena with a shallowly impressed groove running parallel with edge. Thorax (Fig. 3): Black, covered with pollinose vestiture; mesonotum with anterior portion dull, brown; humerus and notopleuron concolorous gray, contrasting distinctly with brown mesonotum, both posterior portions and scutellum shinier, darker, bronzish; scutellum mostly bare, with very few scattered small setae, slightly rounded. Pleural areas entirely pollinose, gray, unicolorous. Femora black but with rather dense covering of gray pollinos- ity, concolorous; apical half of anteroventral edge of fore femur with row of small peglike cuticular bumps best seen in silhouette; tibiae concolorous, mostly pale, tawny but with basal half of dorsal surface invested with silvery-gray pollinosity, mid tibia with larger, ventroapical bristle; hind tibia of some specimens with a long, stout, spinelike, preapical, ventral bristle; basal tarsomeres concolorous with tibial apices, becoming darker apically, apical 2 tarsomeres dark brown. Wing (Fig. 4) entirely hyaline: length-to-width ratio averaging 1:0.45; costal vein ratio averaging 1:0.60: M,.2 vein ratio averaging 1:0.70; posterior crossvein perpendicular with vein M,.»2. Halter pale, whitish-yellow. VOLUME 90, NUMBER 3 561 Figs. 1-3. Eleleides liroceras: 1, Head, frontal view; 2, Same, lateral view; 3, Thorax, dorsal view. 062 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 4. Eleleides liroceras: 4, Wing. Abdomen: Shiny above with metallic dark brown to bronzish-brown re- flections; often with small triangular pollinose gray wedges on lateral margins, variable, many specimens lacking any indication of gray on sides. Sterna 1-6 of female as in Fig. 5; female ventral receptacle as in Figs. 6 and 7. Male genitalia as in generic diagnosis and in Figs. 8, 9, and 10. Type-material—Holotype female, labelled: “ARGENTINA: Province of Buenos Aires, Hurlingham, 26-X-73, D H Habeck, sweeping.” Allotype (in poor condition) and 12 paratypes (all 2 2), with the same label data as the holotype. The holotype (USNM type number 74162), allotype, and 4 para- types were donated to the National Museum of Natural History, Smithsonian Institution, by the Florida State Collection of Arthropods, Gainesville, Florida. The remaining paratypes will be returned to the latter institution. Geographic distribution—Presently known from the type-locality only. Etymology.—Liroceras is a combination of the Greek adjective leiros (Latin transcription liros), meaning “pale” and the noun ceras, meaning “horn, in allusion to the pale third antennal segment. The name stands in apposition to the generic name. Remarks.—The single male specimen was badly damaged although its abdomen was intact, permitting preparation of the genitalic structures for illustration. Eleleides chloris Cresson Figs. 11-13 Eleleides chloris Cresson, 1948:20. Wirth, 1968:142 (review, figures of male genitalia). VOLUME 90, NUMBER 3 563 Figs. 5-10. Eleleides liroceras: 5, Sterna 1-6 of female; 6, Female ventral re- ceptacle; 7, Same; 8, Gonal arch and surstyli of male, lateral view; 9, Surstyli, posterior view; 10, Male genitalia, lateral view. 564 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON aa Figs. 11-13. Eleleides chloris: 11, Male genitalia, lateral view; 12, Aedeagal apodeme and aedeagus, lateral view; 13, Surstyli, posterior view. Diagnosis —Specimens of E. chloris are distinguished by the following combination of characters: eye-to-cheek ratio less than 1:0.25; third an- tennal segment black, appearing pubescent; frons except for cinereous an- terior margin shiny, concolorous with dorsal portion of face, dark bluish green; maxillary palp black; mesonotum more or less uniformly shiny, mostly unicolorous, slightly brassy greenish blue; bristles, especially dorsocentrals, more weakly developed; frequently with one larger pair of acrostichal bristles at level of suture; anterior supra-alar bristle present; femora and tibiae concolorous, dark, but with some surfaces appearing cinereous and lustrous from some angles; abdominal terga unicolorous, dark bluish green, shiny. Male genitalia as in Figs. 11, 12, and 13. Type-material—Holotype female, labelled: “Victoria (Australia) 1888 (hand written)/ 4 /yellowish-orange rectangle/Type (pink, hand written)/ 6678 TYPE Eleleides chloris Cress. (red).” The holotype is in the Academy of Natural Sciences of Philadelphia, type-number 6678. Contrary to what Cresson (1948) cited, the holotype is a female specimen. No other speci- mens of the type-series could be found although Cresson listed 3 male and 2 female paratypes with the same label data as the holotype. VOLUME 90, NUMBER 3 565 Geographic distribution—Southeastern Australia between 30° and 38° south latitude (states of New South Wales, including Australian Capital Territory, and Victoria). Natural history —Wirth (personal communication) collected a series of this species from near the town of Griffith (N.S.W.) in November, 1956, by sweeping emergent vegetation from marshy areas adjacent to a fresh- water creek. Acknowledgments I would like to thank Drs. Willis W. Wirth, F. Christian Thompson, and Oliver S. Flint, Jr., for their critical review and constructive commentary on this paper. I am also appreciative of Dr. Howard V. Weems, Jr. for loaning the specimens of the new species and for permitting the USNM to retain the holotype. Mrs. Elaine R. Hodges prepared Figs. 1-3, Mr. Victor E. Krantz made the wing photo, and Miss Anne Halpern carefully typed the various drafts of this paper. Literature Cited Clausen, P. J. 1973. A revision of the Nearctic species of the genus Pelina (Diptera: Ephydridae). Trans. Amer. Entomol. Soc. 99:119—156. Cresson, E. T., Jr. 1931. Diptera of Patagonia and South Chile. 6(2) Ephydridae. Pp. 85-116. —. 1934. Descriptions of new genera and species of the dipterous family Ephydridae. XI. Trans. Amer. Entomol. Soc. 60:199-222. ——.. 1938. Notes on, and descriptions of, some neotropical Ephydridae (Dipt.). Revta. Entomol. 8:24—40. — —. 1948. A systematic annotated arrangement of the genera and species of the Indo-australian Ephydridae (Diptera). II. The subfamily Notiphilinae and supple- ment to Part I on the subfamily Psilopinae. Trans. Amer. Entomol. Soc. 74: 2s) 1949. FEMUR CLYPEUS ve MAXILLARY PALPUS TIBIA PROBOSCIS iT G IBIAL COMB BASITARSUS “evens Fig. 2. Culicoides furens (Poey), lateral view of female, left wing and right legs removed; for explanation of abbreviations see text. Head (Fig. 3)—Head subspherical to slightly flattened, anterior surface rather flattened and in line with anterior surface of proboscis. Dorsal sur- face of head consisting of broad posterior occiput and undifferentiated an- terior vertex, usually with scattered setae and a row of orbital setae bent over eyes. Eyes bare or with short pubescence between the facets; con- tiguous above bases of antennae, or more or less separated. Two incon- spicuous ocelli located on frons just above bases of antennae. Fronto- clypeus present as a prominent convex sclerite between bases of antennae and proboscis. VOLUME 90, NUMBER 3 619 VERT oe z MEDIAN BRISTLE BASE------4 OeEERUseee eS : COMPOUND EYE --~~--- af FRONTO-CLYPEUS _____. OL RNCTORYs a uilrs Lf | \ SS \ | Zé, / \ A NX \ | SSH AROSE / \\” LABRUM-EP I PHARYNX ae LACINIA / es LATERAL TEETH \ / eS HYPOPHARYNX “.LABELLUM \----TERMINAL TEETH A, FEMALE HEAD OTHESIERNUMMe se 2 CAUDOMEDIAN EXCAVATION —--~~ BASISTYLE —----—-—---- ~------ PARAMERE ~-----+------------- AEDEAGUS Di STT STVILB 2 SS SS SISO SSS QTH TERGUM APICOLATERAL PROCESS PSUS GS SS SSS SS aS SSS a= CERCUS B, MALE GENITALIA Some) Fig. 3. Culicoides ssp., parts labeled. a, Female head; b, Male genitalia (after Amaud, 1956). Antenna usually 15-segmented, but basal segment or scape reduced to a ringlike segment in head capsule hidden by the enlarged pedicel (in using the keys, the presence of the scape is always implied and included in the count of segments). The 13 divisions of the flagellum are not true 620 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON morphological segments, but in this series of papers they are counted as segments in accordance with established custom. First 8 segments of fe- male antenna similar and distinct from last 5, which are usually more elongated. Antennal ratio (AR) is the ratio of combined lengths of last five segments to the preceding 8. Antennal segments 4-10 of female usually short and typically bearing basal ring of long verticils and subapical pair of hyaline sensory setae; last 5 usually elongate with basal verticils not so well developed, and usually with scattered short setae and sensory setae on distal portion. Male antennae more or less plumose, pedicel much larger than in female, first 9 or 10 flagellar segments short and similar, bear- ing oblique whorls of long verticils, and distal 3 or 4 segments much elongated, with short verticils. Mouthparts well developed, stronger in females than in males, fitted for piercing and sucking, and forming an elongate proboscis often as long as head. Labium fleshy with scattered setae, enclosing the other 6 parts, which are slender, distally toothed blades of subequal length. These con- sisting of a strong labrum, a pair of mandibles, a pair of maxillae, and a tubular sclerotized hypopharynx. Maxillary palpi usually 5-segmented in both sexes, setose; third segment usually with distal sensory organ and often swollen. Thorax.—Usually rather broad and convex above, arched anteriorly but not greatly projecting over head, prescutellar area flattened. Pronotum divided into lateral halves by anterior development of mesonotum, partly hidden just below humeri. Mesonotum sometimes bearing small spine or tubercle in middle of anterior margin; a pair of small humeral pits located laterally near anterior margin. Pseudosutural foveae extending from the pits caudad to near ends of scutellum as a sublateral pair of slightly depressed lines. Lateral paratergites well developed in the higher Ceratopogoninae. Scutellum prominent, transverse, usually with bristles; postscutellum arched, bare, with small deep central glandular depression, but without longi- tudinal division. Legs.—Legs usually rather short and stout, with characteristic modifi- cations useful in generic differentiation. One or more pairs of femora may be swollen and bearing prominent ventral spines; tarsomeres may bear spines, or shapes of fourth or fifth tarsomeres may be characteristic. Presence of long, blunt, black, non-tapering spines or “batonnets” on ven- tral side of fifth tarsomere characteristic of some genera. Shape and de- velopment of the claws are extremely important in generic separation, and presence of a well-developed empodium is used to characterize the For- cipomyiinae. The most useful modifications of the fifth tarsomeres and claws are found only in the female sex, and thus the males are more difficult to characterize generically without dissecting the genitalia. Hind tarsal VOLUME 90, NUMBER 3 621 ratio (abbreviated TR), often useful in specific and generic diagnoses, ob- tained by dividing length of basitarsus by length of second tarsomere. Wing.—Wing moderately broad (narrower in male) with apex evenly rounded, anal lobe rarely prominent, alula usually without fringe. Wing venation extremely important in generic classification. The Tillyard modi- fication of the Comstock-Needham system of venation is followed here (see Fig. 2). The most prominent general features are reduction of the radius to a compact anterior system, fusion of R2 + 3 with R4 + 5 to form radial sector (Rs), termination of Rl and Rs in the costa, and forking of the an- terior media and its union to the radius by a crossvein. Costa long or short; one or two anterior radial cells present, one or other long or short, reduced, or absent; fork of anterior media sessile, petiolate, or interrupted beyond the crossvein. The costal ratio (CR) is obtained by dividing length of costa by wing length, measuring both from the level of the basal arculus of the wing. Posterior veins (M1, M2, M3 + 4, Cul, Cu2, and Ist A) relatively weak. Development or reduction of the macrotrichia (long hairs) or micro- trichia (microscopic spicules or setae) and presence or absence of a pattern of pigmented spots or markings are useful. In life, the wings are almost always superimposed and folded flat over the back while the insect is at rest, thus differing in habits from non-biting midges of the family Chirono- midae that never superimpose their wings over the back. Abdomen.—Female abdomen usually short and blunt, somewhat dorso- ventrally flattened and terminating in a pair of cerci which are short and rounded or rarely (in Leptoconops) long and tapered. Genital opening in some genera flanked by modifications of the sterna or an armature of spines or hairs. Spermathecae usually strongly sclerotized, their number, size, and shape very useful in classification. In the genera of the Palpomyi- ini and Stenoxenini paired eversible glands open on anterior margins of certain terga; these openings marked internally by tapered internal sclero- tized apodemes (“gland rods”) extending from bases of terga toward base of abdomen. Male genitalia (Fig. 3)—Male genitalia of simple nematocerous type. Rotation of eighth and succeeding segments occurring to greater or lesser de- gree. Ninth tergum usually an expanded plate, convex externally and hollowed out mesad, and bearing the anus flanked by pair of membranous cerci on ventromesal face. Ninth sternum usually much shorter and trans- verse, with aedeagus projecting caudoventrad from posterior margin and articulating with bases of basistyles. Genital appendages or claspers (par- ameres of Snodgrass, 1957) prominent, forcepslike and 2-segmented; basi- styles usually fairly large and simple, bearing a more or less tapered, usu- ally simple, incurved dististyle. Aedeagus usually more or less triangular or Y-shaped in ventral view, flattened and bent in gutter shape in cross 622 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON section, convex ventrally and bearing the membranous penis dorsally (internally) with its opening at or near tip of aedeagus. Parameres (claspettes of Snodgrass) typically a pair of submedian processes, often fused, usually symmetrical, articulating with bases of basistyles on dorsal side and pro- jecting distad between aedeagus and ventral face of ninth tergum. IV. Biology and Immature Stages of Ceratopogonidae The first important studies on the immature stages of biting midges were made by Gercke (1883), Meinert (1886), and Mik (1888) in Europe, and Long (1902) and Johannsen (1905) in North America. Later the European workers Rieth (1915), Goetghebuer (1920), Saunders (1924), Kieffer (1925), Thienemann (1928), Lenz (1934), and Mayer (1934) brought knowledge of the European species up to date, and Malloch (1914, 1915, 1917) and Thomsen (1937) reviewed the North American species. Relatively little | comprehensive work has been done in other regions of the world, although many scattered descriptions have appeared. Hennig (1950) reviewed all the published information on immature stages up to that date. Wirth (1952) published what was known of the species occurring in California, and essentially the same information was summarized in Wirth and Stone (1963). Serious comprehensive work on the immature stages has been done only in the genera Forcipomyia (Saunders, 1924, 1925, 1956, 1959; Chan and LeRoux, 1965, 1970, 197la, b, c); Atrichopogon (Nielsen, 1951; Ewen and Saunders, 1958); and Culicoides (Lawson, 1951; Kettle and Lawson, 1953; and Jamnback, 1965). Immature stages of the Forcipomyiinae are especially useful in classification and separation of closely related species, and reflect important ecological adaptations. Egg.—kEggs differing markedly among subfamilies (Figs. 3-7); oval to elongate, often black and glossy (Leptoconopinae and Forcipomyiinae), U-shaped and enclosed in gelatinous cover in Dasyheleinae, and elongate and banana shaped in Ceratopogoninae, with conspicuous frilled collar on one end in Stenoxenini. They may be laid on a moist substrate in loose groups (Forcipomyia, Culicoides, and most other genera), or in strings or masses with gelatinous coating (Bezzia, Mallochohelea, etc.). Larva.—Without functional spiracles. Head sclerotized (except Lepto- conopinae), with strong, toothed, non-opposable mandibles; internally with conspicuous pharyngeal apparatus consisting of two strongly diverging arms and a series of combs serving to reduce and sort the food. A short collar between head and thorax; 3 thoracic and 9 abdominal segments. Four larval instars. General habitus and structure of the larva differs greatly among the subfamilies as follows: Leptoconopinae (Fig. 4): Larva elongate and smooth, without con- VOLUME 90, NUMBER 3 623 Fig. 4. Life stages of Leptoconopinae: a-f, Leptoconops spinosifrons (Carter) (after Laurence and Mathias); g, Leptoconops americanus (drawn by Ethel Grogan). a, Larva; b, Larval head; c, Pupa; d, Larval pharyngeal apparatus; e, Larval mandible; f, Egg. spicuous setae, many segments subdivided into two subsegments; anal segment with three short lobes. Head prognathous, head capsule not fully sclerotized, with internal rodlike structures of uncertain morphology. Larval biology adapted to arid or desert areas and coastal or inland beaches in moist or wet soil which is usually hypersaline; the larvae burrow in the soil, often 624 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 5. Life stages of Forcipomyiinae (Forcipomyia ingrami Carter) (after Williams, 1944). a, Pupa; b, Adult male; c, Distal antennal segment of female; d, Male genitalia; e, Egg cluster; f, Larval mandible; g, Larva. to great depths following moisture downward as the soil dries and cracks; some species aestivate during extremely dry periods. Forcipomyiinae (Fig. 5): Larva often relatively stout; head usually hypognathous; body segments with conspicuous setae, often complex and set on projecting tubercles; prothorax with bilobed anterior proleg, last segment VOLUME 90, NUMBER 3 vn ~~ yy ‘hay GU SS xX Nniiy nay NN \ \ \ ( Fig. 6. Life stages of Dasyheleinae: hawaiiensis Macfie) (after Williams, 1944). d, Caudal end of larva; e, h, Egg cluster; f Cyiaae . 7 \, NL a Sy ‘ ’ y Volare . 7. “2 , on \) CR ab iat q bn a i 24) 3 “s § a she f af sign tyne isc raf - #3 y WS ) , N go K BER . 4 > =e Aarh 1. Ht FH hy Aas —Fag Sebi 4 ah 4 ° of *4 q fe ¥ oy i "4 Wy = ory Ap ae d = saad ae: = a —— 625 (a-e, Dasyhelea calvescens Mactie; t-i, D. a, Adult female; b, g, Pupa; ce ~ Larva; i, Larval mandible. _ Male wing: 626 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON yp ier at y | A Pei Z Wp Wp Fig. 7. Life stages of Ceratopogoninae, tribe Culicoidini: a, Culicoides guttipennis (Coquillett), adult female (drawn by Ethel Grogan); b-c, C. accraensis Carter, Ingram and Macfie (after Carter, Ingram and Macfie, 1921). b, Pupa; c, Larva, details of larval head, caudal segment. VOLUME 90, NUMBER 3 627 with posterior proleg bearing conspicuous hooklets. Habits crawling; usu- ally terrestrial in damp places such as under bark or in mosses, or semi- aquatic; feeding on fungi, algae, or plant debris. In some genera last larval skin retained over posterior segments of pupa. Dasyheleinae (Fig. 6): Larva moderately elongate, head hypognathous or intermediate in shape; body without conspicuous hairs or projections; an- terior proleg absent, posterior proleg present with hooklets much as in Forcipomyiinae. Habits wriggling chironomid-like, not free swimming; usu- ally living in blanket algae or algae on mud or wet rocks at stream or pond margins; often in restricted habitats such as rock pools, tree holes and sap flows. Feeding habits usually phytophagous on algae. Ceratopogoninae (Fig. 7): Larva with distinctive, elongate, eellike form and prognathous, somewhat elongated head. Body hairs minute except on last segment; cuticle smooth, whitish or translucent; no prolegs (except first instar of some Culicoides). Habits ranging from semiaquatic, burrow- ing in moist soil or sphagnum mats, to fully aquatic and free swimming; some genera in benthos and plankton of large lakes and streams. Feeding habits usually carnivorous. Pupa.—Appearance relatively uniform and _ characteristic; yellowish brown to blackish in color; usually rather conical in form, body more or less compact, stout anteriorly with pair of prothoracic respiratory horns of diverse structure; last segment bearing pair of pointed, apicolateral pro- cesses. Structural details (Figs. 4-7) of respiratory horn, operculum (vertex), tubercles bearing spines or setae, and apicolateral processes of abdomen usually of great diagnostic value for taxa at all levels. Movement usually possible by twisting movements of abdomen but not true swimming; air breathing by anterior horns, aquatic forms hanging at water surface by non-wettable horns, supported by air bubble beneath wing cases. V. Systematic Arrangement of the North American Genera of Ceratopogonidae Subfamily Leptoconopinae Noé Genus Leptoconops Skuse (23 North American species) Subfamily Forcipomyiinae Lenz Genus Atrichopogon Kieffer (27 species) Genus Forcipomyia Meigen (73 species) Subfamily Dasyheleinae Lenz Genus Dasyhelea Kieffer (29 species) 628 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Subfamily Ceratopogoninae Newman Tribe Culicoidini Kieffer Genus Paradasyhelea Macfie (1 species) Genus Culicoides Latreille (137 species) Tribe Ceratopogonini Newman Genus Alluaudomyia Kieffer (8 species) Genus Brachypogon Kieffer (1 species) Genus Ceratoculicoides Wirth and Ratanaworabhan (3 species) Genus Ceratopogon Meigen (4 species) Genus Isohelea Kieffer (3 species) Genus Rhynchohelea Wirth and Blanton (1 species) Tribe Stilobezziini Wirth Genus Echinohelea Macfie (1 species) Genus Monohelea Kieffer (16 species) Genus Parabezzia Malloch (14 species) Genus Serromyia Meigen (3 species) Genus Stilobezzia Kieffer (18 species) Tribe Heteromyiini Wirth Genus Clinohelea Kieffer (7 species) Genus Heteromyia Say (2 species) Genus Neurobezzia Wirth and Ratanaworabhan (1 species) Genus Neurohelea Kieffer (2 species) Genus Pellucidomyia Mactie (1 species) Tribe Sphaeromiini Newman Genus Jenkinshelea Mactie (4 species) Genus Johannsenomyia Malloch (2 species) Genus Macropeza Meigen (1 species) Genus Mallochohelea Wirth (11 species) Genus Nilobezzia Kieffer (4 species) Genus Probezzia Kieffer (19 species) Genus Sphaeromias Curtis (1 species) Tribe Palpomyiini Enderlein Genus Bezzia Kieffer (43 species) Genus Pachyhelea Wirth (1 species) Genus Palpomyia Meigen (25 species) VOLUME 90, NUMBER 3 629 Tribe Stenoxenini Coquillett Genus Paryphoconus Enderlein (1 species) Genus Stenoxenus Coquillett (1 species) VI. General Classification of the Ceratopogonidae Leptoconopinae.—There is general agreement that the Leptoconopinae stand somewhat apart from the other ceratopogonids and some modern workers have proposed separate family status for them (Zilahi-Sebess, 1960; Krivosheina, 1962; Remm, 1975). The wide separation of the eyes in both sexes, 13- or 14-segmented female antennae, pale wings without macrotrichia, absence of the r-m crossvein and extreme shortening of the radial system into a compact stigma, lengthening of the female cerci into tapering lamellae in most, articulated tooth at the tip of the male dististyle, and extreme reduction of the larval head capsule are unusual in the Cera- topogonidae. Great as these differences are, we do not believe the Lepto- conopinae are distinctive enough from other ceratopogonids to warrant family recognition. Forcipomyiinae.—Classification of the Forcipomyiinae is more advanced than that of any other group, thanks almost entirely to fine work on the immature stages by Saunders. Chan and LeRoux (1965, 1970, 1971a, b, c) have continued more recently in Saunders’ tradition in describing several new subgenera of Forcipomyia, emphasizing all the while the importance of immature stages in the classification of this subfamily. Dessart (1963), working on cacao pollination in Zaire, brought the African species into the modern classification. Remm (196la, b, 1962, 1969, 1974a, b), in Estonia, has made similar advances in revising the Palaearctic species that are found in the Soviet Republics. Larvae, pupae, male genitalia, and habits offer excellent characters in the Forcipomyiinae, but the females are more diffi- cult to classify. Dasyheleinae—In some ways the genus Dasyhelea is as distinct as Leptoconops, although in general its characters are intermediate between those of the Forcipomyiinae and Ceratopogoninae, as the older workers recognized when they proposed the term “Ceratopogoinae Intermediae” for these midges (Rieth, 1915). The distinctive sculpturing of the antennal segments, the characteristic flattening of the head with closely contiguous eyes and reduced mouthparts not adapted for bloodsucking in the female, the asymmetrical male parameres, and the wriggling type of larva with only the posterior proleg present, serve to place this genus in a subfamily of its Own. Ceratopogoninae.—We recognize seven tribes in this subfamily, as listed in our synopsis. The genera placed in the tribe Culicoidini appear to be as primitive and nonspecialized as any in the family, and along with those in 630 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON the subfamilies Dasyheleinae and Leptoconopinae and the tribe Ceratopo- gonini may give us more clues to the ancestral lineage than other sections of the family. We believe it is significant that the annectant genera Para- dasyhelea Macfie and Austroconops Wirth and Lee from Australia, New Zealand, and Patagonia belong in the Culicoidini. Classification of the important blood-sucking genus Culicoides is still poorly understood, for although many subgenera have been proposed, most of them are valid only in one particular geographic region or another and must be revised or supplemented to bring the other species of the world into the system. The tribe Ceratopogonini has been relatively neglected and needs a good contemporary revisionary study. Distinctions between this tribe and the Stilobezziini are not clear-cut and good synapomorphies have not been worked out in either tribe. Classification of these tribes, as for others in the Ceratopogoninae, has been seriously hindered by lack of good characters in the immature stages. Generic characters have been found mainly in the female wings, fifth tarsomeres and claws, and the male genitalia. Tokunaga (1962) in his revision of the New Guinea “Palpomyiinae” adopted a conservative viewpoint with respect to these large predaceous midges, us- ing the traditional characters of number of radial cells in the wing and armature of the femora. A much better classification of genera can be secured, however, by first separating those whose females have internal sclerotized gland rods and eversible glands on the abdomen. These genera, comprising the tribes Stenoxenini and Palpomyiini, are also characterized by the females having cordiform fourth tarsomeres, fifth tarsomeres with- out true batonnets, and claws simple and equal. Wirth (1962) published this separation in 1962, when he gave a key and suggested an arrange- ment for the remaining genera, which he placed in two tribes, the Hetero- myiini and Sphaeromiini. In the tribe Heteromyiini, the female claws are unequal, at least on the hind pair of legs; the fifth tarsomere does not bear ventral batonnets and those on the fore legs are often inflated, while the fourth tarsomere is sometimes divided into spinose bifid lobes. Larger genera of the tribe Heteromyiini are Clinohelea Kieffer and Neurohelea Kieffer, and in the tropics are such bizarre forms as Heteromyia Say, Pellucidomyia Macfie, and Tetrabezzia Kieffer. In the genera of the tribe Sphaeromiini are found the most striking developments of the secondary sexual characters of the female tarsi. Com- mon to all these genera is the development of specialized, stout, blunt, black, ventral spines on the fifth tarsomere which Kieffer called “batonnets.” Among genera, these are arranged differently. Other convenient charac- ters used for generic separation are the equal or unequal size of the fe- male claws, location of the basal tooth on the internal or external side of VOLUME 90, NUMBER 3 631 each claw, and the shape of the claw, straight, bent, or curved. The length of the costa is a more valuable character than the presence of one or two radial cells, and the armature of the femur is not often useful above the species level. In addition, certain genera characteristically have dull polli- nose integument, especially on the thorax, whereas others are shining midges. In the tribe Sphaeromiini, the pupae of some genera have unusual ventral glandular discs on certain abdominal segments which appear to be used to attach the pupa fast to emergent vegetation or other objects above the water level preparatory to eclosion. This development probably is associated with the preferred larval habitat, larger streams with fluctuat- ing water level or lakes with considerable wave action on the shores. VII. Key to North American Genera of Ceratopogonidae (Adults, Primarily for Females) North American genera not occurring in the Washington, D.C. area are bracketed in this key. I Wing with crossvein r-m absent (Fig. 8); wing without macro- trichia (Fig. 8); female antenna with 13 or 14 segments (Fig. 14, flagellum) (LEPTOCONOPINAE) [Leptoconops Kieffer] — Wing with crossvein r-m present (Fig. 9); wing with (Fig. 10) or without macrotrichia; female antenna with 15 segments (Fig. 15, flagellum) We Empodium well developed, at least in female, claws markedly curved (Figs. 18, 20); wing usually with numerous macro- trichia (Figs. 9-10) (FORCIPOMYIINAE) 3 ~ Empodium smaller or vestigal (Fig. 24); claws usually gently curved (Fig. 48) but in some females of specialized raptorial form (Fig. 70); wing usually with microtrichia less numerous or absent 4 3. Costa reaching well beyond middle of wing; second radial cell longer than first and usually broad; microtrichia of wing large and conspicuous, macrotrichia when present scattered, suberect, not scalelike; fringe of posterior border of wing simple, a single row of alternating short and very short, simple, straight hairs (Fig. 9) Atrichopogon Kietter ~ Costa short or long; second radial cell if long is narrow; micro- trichia minute, macrotrichia more abundant, covering greater part of wing, sloping, often scalelike; fringe more complex, not bo a single row of setae (Fig. 10) Forcipomyia Meigen 4, Antennal segments sculptured, at least in male (Fig. 34); first radial cell nearly or completely closed, second closed or square 10. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON ended, usually ending near middle of wing (Fig. 29); female claws small and equal (Fig. 48); eyes very finely pubescent | (DASYHELEINAE) Dasyhelea Kieffer Antennal segments not sculptured; one or both radial cells well developed, second not markedly square ended, usually ending past middle of wing (except in Paradasyhelea); eyes usually bare (CERATOPOGONINAE) 3D Media petiolate, forking distad of level of r-m crossvein (Fig. 77) (in Echinohelea, with spinose legs, media forks just at crossvein (Fig. 81)); M2 sometimes obsolescent basally 6 Media sessile, forking at or proximad of level of crossvein (Fig. 147); M2 nearly always complete 19 Claws of both sexes small, equal and simple (Fig. 45); macro- trichia of wing usually abundant; two more or less equal radial cells usually present; humeral pits prominent (CULI- COIDINIT) if Claws of female usually large, equal or unequal (Fig. 68), those of male smaller and equal; macrotrichia less numerous or absent; one or two radial cells, second usually larger than first (Fig. 78); humeral pits less prominent or absent 8 Two radial cells present, usually well formed, rarely obliterated (Fig. 31); costa usually extending past middle of wing; wing usually adorned with pale or dark spots (Fig. 31); palpus 5- segmented (Fig. 40); female mouthparts usually fitted for bloodsucking with toothed mandibles (Fig. 38) Culicoides Latreille Both radial cells completely obliterated, costa short, not reaching middle of wing; wing not conspicuously adorned (Fig. 30); palpus 3- or 4-segmented; mouthparts reduced, female mandibles not toothed [Paradasyhelea Macfie] Second radial cell small, not or little longer than first, one or both radial cells may be closed (in Alluaudomyia first radial cell closed, second long, macrotrichia usually numerous, wing usually with small black spots (Fig. 53)); eyes usually pubes- cent; wing often milky (CERATOPOGONINI) 9 Second radial cell long, much longer than first, which may be obliterated (Fig. 77); wing hyaline or with dark pattern, not milky; eyes usually bare (STILOBEZZIINI) 14 Wing with both radial cells open and complete (Fig. 56) 10 Wing with at least one radial cell obsolete (Fig. 57) 11 Costa extending to well past middle of wing, radial cells more elongate (Fig. 55); three large spermathecae present (Fig. 72); male parameres separate (Fig. 74) Ceratopogon Meigen VOLUME 90, NUMBER 3 633 Jig 12. 13. 16. les Costa extending to about middle of wing; radial cells short with adjacent veins thickened (Fig. 56); one or two large spermathecae present; male parameres usually broadly fused proximally Isohelea Keitfer Female antenna 14-segmented, very short, with segments moniliform (Fig. 64); proboscis stout and truncate (Fig. 59); palpus stout, third segment greatly broadened (Fig. 61); radial cells absent, vein M1 obsolete distally, M2 absent (Fig. 58) [Rhynchohelea Wirth and Blanton] Female antenna 15-segmented, elongate with slender segments (Fig. 66); radial cells and media various YZ Female claws unequal, at least on hind legs (Fig. 68); wing pattern of small isolated black spots or streaks (Fig. 53) Alluaudomyia Kieffer Female claws equal on all legs; wing pattern present or ab- sent 13 Female claws small to large, but of same length on all legs (Fig. 70); one spermatheca; eyes contiguous; radial cells obsolete; male apicolateral processes absent Brachypogon Kiefter Female claws small on one pair of legs, usually hind legs (Fig. 67); 2 spermathecae; eyes broadly separated; first radial cell closed, second present but small (Fig. 57); male ninth tergum with well developed apicolateral processes Ceratoculicoides Wirth and Ratanaworabhan Femora armed with one or more stout ventral spines, at least on one pair of legs (Figs. 91, 94) 15 Femora not armed with stout ventral spines 17 All femora armed with numerous spines, at least in male sex, these not confined ventrally (Fig. 94); two well developed ra- dial cells (Fig. 81); male antenna not plumose, last five seg- ments elongated as in female; reddish yellow pruinose spe- cies Echinohelea Mactie Only hind or fore femur armed with ventral spines; one or two radial cells; male antenna plumose; color various 16 Fore femur slender and unarmed, hind femur greatly swollen and arcuate, and armed with numerous ventral spines (Fig. 91); two well developed radial cells (Fig. 80); shining black species Serromyia Meigen Fore femur armed with 1-2 ventral spines; hind femur slender and unarmed; first radial cell obliterated (Fig. 77); pollinose yellow or brownish species Stilobezzia subgenus Eukraiohelea Ingram and \lactie One radial cell present; female costa long, extending past end 634 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON of vein R4 + 5 and ending nearly at wing tip (Fig. 82); palpus with segments four and five fused (Fig. 86) | Parabezzia Malloch Two radial cells present; costa shorter, not extending past end of vein R4 + 5 and not ending nearly at wing tip (Fig. 78); palpus 5-segmented (Fig. 87) 18 Female claws usually large and unequal on all legs (Fig. 95) Stilobezzia Kietter Female claws equal on four anterior legs, hind leg with one long talon and with or without a second or shorter claw Monohelea Kieffer Fifth tarsomeres of female armed ventrally with stout, black, blunt spines (batonnets) (Fig. 138); female abdomen without internal sclerotized gland rods; eighth segment of female often with ventral pair of hair tufts near gonopore (Fig. 141) (SPHAEROMIINT) 20 Fifth tarsomeres of female unarmed or provided with only slender, sharp-tipped spines (Fig. 121); if tarsi armed, female abdomen with internal sclerotized gland rods (Fig. 162); eighth segment of female without ventral hair tufts near gonopore 26 Female tarsal claws unequal on four posterior legs, equal on fore legs (Fig. 134); femora unarmed (abdomen petiolate; eighth segment of female abdomen with ventral pair of hair tufts; mesonotum with microtubercles bearing appressed setae (Fig. 132); costa extending to about 0.8 of wing length; usu- ally two radial cells (Fig. 123)). Johannsenomyia Malloch Female tarsal claws equal on all legs; femora armed or un- armed Ok Female claws with slender internal basal barb and gently curved distally (Fig. 137); costa extending nearly to wing tao) (liikes, 125) Sphaeromias Kieffer Female claws with blunt external basal tooth at least on one pair of legs and usually straight or flattened distally (Fig. 138); costa various 22 Costa short, extending to about 0.8 of wing length (Fig. 122) 23 Costa long, extending nearly to wing tip (CR over 0.87) (Fig. 126) 24 Body slender; mesonotum shining yellow to black with little or no pollen; femora armed or unarmed; two radial cells present (Fig. 122); male genitalia with well developed basi- style and articulated dististyle (Fig. 142) Mallochohelea Wirth Body stout; mesonotum dull, usually with dense whitish to grayish pollen; femora ventrally and tibiae dorsally armed ee VOLUME 90, NUMBER 3 63 24. 25. 26(19). 2. 28. 29. 30. 31(26). Ol with numerous fine sharp spines; one or two radial cells present (Fig. 124); male genitalia with short, stout basistyle fused with budlike dististyle (Fig. 140) Nilobezzia Kieffer Wing broad with large angular anal lobe (Fig. 128) Jenkinshelea Macfie Wing not unusually broad; anal lobe not angularly developed 25 Female claws short and curved, sharp pointed, each with in- conspicuous external basal tooth (Fig. 145); mesonotum without strong erect bristles [Macropeza Meigen] Female claws longer, straighter, and somewhat flattened dis- tally, each with strong, blunt, external basal tooth; meso- notum with strong erect bristles Probezzia Kieffer Claws of female usually unequal (at least on hind leg), or a single claw with basal tooth (Figs. 112, 114); female abdomen without internal sclerotized gland rods (HETEROMYIINT) Pat Claws of female equal on all legs (Fig. 159); female abdomen usually with one or more pairs of internal sclerotized gland rods (Fig. 162) 31 Costa greatly prolonged beyond tip of vein Rs (Figs. 100- 101) 28 Costa not prolonged beyond tip of vein Rs (Figs. 102-104) 29 Two radial cells present (Fig. 101); claws equal on all legs; fifth tarsomere of fore leg somewhat inflated (Fig. 111) Neurohelea Kiettfer One radial cell present (Fig. 100); claws equal on fore and mid legs, unequal on hind leg; fifth tarsomere of fore leg not inflated (Fig. 113) [Neurobezzia Wirth and Ratanaworabhan | Fourth tarsomere cylindrical or cordiform but not divided into spinose bifid lobes; claws equal on fore and mid legs, with one long talon and much smaller second claw (Fig. 114) 30 Fourth tarsomere of mid and hind legs ending in two bifid lobes armed with spines, cordiform on fore leg; claws equal on fore leg, unequal on mid and hind legs (Fig. 112) Clinohelea Kietter Fore femur greatly swollen and armed ventrally, the tibia arcuate (Fig. 110); fourth tarsomeres of fore and mid legs cordate (Fig. 114); one or two radial cells; wing fasciate (Fig. 103); mesonotum more or less shining Heteromyia Say Fore femur slender, unarmed, the tibia normal; fourth tarso- meres not cordate; one radial cell (Fig. 102); wing milky white, mesonotum whitish pollinose [Pellucidomyia Macfie} Body unusually slender and dorsoventrally flattened; one very narrow radial cell usually extending to wing tip (Fig. 150); r-m crossvein often very short, cell between base of R and 636 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON M very narrow or even obliterated (Fig. 151); eyes broadly separated (Fig. 152); eighth abdominal segment of female narrow distally with genital sclerotization; legs long and slender, with fine setae; femora unarmed, claws very short (Fig. 160) (STENOXENINI) on — Body not unusually slender or dorsoventrally flattened; one or two radial cells, if cells narrow costa not extending nearly to wing tip; r-m crossvein longer, cell between R and M well formed and not very narrow (Figs. 147-149); eyes narrowly to moderately separated; eighth abdominal segment of female not modified; legs usually not extremely long nor hairy; femora often armed; claws usually moderately long (Figs. 159-161) (PALPOMYIINIT) 33 32, Thorax broadly rounded anteriorly, without median spine; fe- male wing with vein M2 strikingly elbowed at base (Fig. 151); palpus 4-segmented; two spermathecae present; male para- meres fused apically [Stenoxenus Coquillett ] = Thorax narrowed in front, more or less conical with erect an- teromedian spine (Fig. 154); female with vein M2 not elbowed at base (Fig. 150); palpus 5-segmented; one spermatheca pres- ent; male parameres separate apically — [Paryphoconus Enderlein| 33: Two radial cells 34 — One radial cell (Fig. 149) Bezzia Kieffer 34. Hind femur greatly swollen; femora unarmed (Fig. 158) [Pachyhelea Wirth] - Hind femur not greatly swollen, if moderately swollen at least one pair of femora armed with ventral spines; fore femur often swollen (Fig. 157) Palpomyia Meigen Acknowledgments We gratefully acknowledge support of this project in the form of a re- search grant from the Washington Biologists Field Club. We especially thank C. Gordon Fredine, President, and J. F. G. Clarke, Chairman of the Research Committee, for the year 1975-76. Mr. William L. Grogan, Jr. of the University of Maryland, College Park, carried on the field work on Plummers Island and environs with financial support from this grant during the summer of 1976. For permission to collect in the Chesapeake and Ohio National Historical Park we wish to thank William R. Tailor, Superintendent. We thank William H. Stickel, U.S. Fish and Wildlife Service, Laurel, Maryland, for permission to collect in the Patuxent Wild- life Research Center. We also are grateful to Mrs. Ethel L. Grogan for assistance with the illustrations. VOLUME 90, NUMBER 3 637 => Figs. 8-28. Chracters of Leptoconopinae and Forcipomyiinae (Figs. 8-164 drawn by N. C. Ratanaworabhan). Leptoconops americanus Carter: 8, 11, 13, 14, 16, 19, 24. 28. Forcipomyia bipunctata (Linnaeus): 10, 12, 15, 17, 18, 21, 22, 23, 26. Forcipomyia glauca Macfie: 25. Atrichopogon levis (Coquillett): 9, 20, 27. Figs. 29-52. Characters of Dasyheleinae and Ceratopogoninae (Culicoidini). Dasyhelea grisea (Coquillett): 32, 34, 42, 46, 48, 51. Dasyhelea mutabilis (Coquillett): 29. Paradasyhelea minuta Wirth and Lee: 30, 36, 37, 39, 43, 47, 52. Culicoides yukonensis Hoffman: 31, 33, 35, 38, 40, 41, 44, 45, 49, 50. Figs. 53-76. Characters of Ceratopogoninae (Ceratopogonini). Alluaudomyia bella (Coquillett): 53, 68, 73, 76. Brachypogon paraensis Wirth and Blanton: 54, 60, 63, 66, 70, 71, 75. Ceratoculicoides longipennis Wirth: 57, 62, 67. Ceratopogon culicoidithorax Hoffman: 55, 69, 72, 74. Isohelea stigmalis (Coquillett): 56, 65. Rhynchohelea monilicornis Wirth and Blanton: 58, 59, 64. Figs. 77-99. Characters of Ceratopogoninae (Stilobezziini). Stilobezzia elegantula (Johannsen): 77. Stilobezzia festiva Kieffer: 78, 88, 90, 95. Echinohelea lanei Wirth: 81, 83, 92, 94, 96, 97. Monohelea hieroglyphica Kieffer: 79, 87, 89, 93, 98, 99. Parabezzia petiolata Malloch: 82, 85, 86. Serromyia femorata (Meigen): 80, 84, 91. Figs. 100-121. Characters of Ceratopogoniae (Heteromyiini). Clinohelea bimaculata (Loew): 104, 109, 112, 119, 120. Heteromyia fasciata (Say): 103, 105, 106, 107, 108, 110, 114. Pellucidomyia lanei Wirth: 102, 115, 116. Neurobezzia granulosa (Wirth): 100, 113, 121. Neurohelea nigra Wirth: 101, 111, 117, 118. Figs. 122-146. Characters of Ceratopogoninae (Sphaeromiini). Jenkinshelea magni- pennis (Johannsen): 128, 135. Johannsenomyia argentata (Loew): 123, 129, 132, 133, 134, 136, 144, 146. Macropeza natalensis (de Meillon): 127, 130, 145. Mallocho- helea albibasis (Malloch): 122, 142, 143. Nilobezzia schwarzii (Coquillett): 124, 138, 140. Probezzia pallida Malloch: 126, 139, 141. Sphaeromias longipennis (Loew): 125, WSC, Le Figs. 147-164. Characters of Ceratopogoninae (Palpomyiini and Stenoxenini). Bezzia setulosa (Loew): 149, 153, 163, 164. Pachyhelea pachymera (Williston): 148, 158, 161. Palpomyia plebeia (Loew): 147, 155, 157, 159, 162. Stenoxenus coomani Seguy: 151, 152, 156, 160. Paryphoconus angustipennis Enderlein: 150, 154. 638 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON pir SSS aT i Tj Ma TL 10 A y) Ha \! He} hint i a oe a 639 VOLUME 90, NUMBER 3 ace ae = Gy LO = =e = _-~™ ee ee I — 640 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON : {> LUO O000 COOL» ; Aft AEH LIE LAO ae ye age aes are SN ies ree gar cgeeee atin arsed ean a = O SQ Gna Gn a Cams Oo oO MAM pd OO COO EE re IES Re ees 71 VOLUME 90, NUMBER 3 641 in ie iin 2 ee Lat } Hi I {h IMMA cone aah anainnyarieaee | 5 PA ™, Ve Ne Se MMWiteiy epimers AT? gene i 7 8 KA Ah boll MeO EE re a { yg ee a ee pee Ta : a Hesbeon cea eee wd Deere Fe [4 kA 2 Aina : ; LM Ot enna x ae \ i ~ A ; ~y a : ‘ iS L 95 & |r x \ ; ~N s \\ . \ — PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 642 x es MLE aoe . 2 6-9 spinules, lateral margin unarmed; dorsomesial distal angle with strong | spine, mesial margin with long setae. First segment with lateral margin unarmed or with small spinule; ventral margin produced, unarmed. An- tennal acicle as long as, or a little shorter than ocular peduncles; arcuate, subtriangular, with dorsal surface flattened; mesial and lateral margins — unarmed, mesial margin with long setae. Antennal flagella long, over- reaching right cheliped, articles with extremely short setae. Mandible without distinctive characters. Maxillule with proximal endite subquadrate, internal lobe of endopodite well developed and with one bristle, external lobe produced, ovoid, not recurved. Maxilla with endop- odite inflated basally, tapering to tip, exceeding scaphognathite in distal extension. First maxilliped with endopodite about half length of exopodite; basal portion of exopodite very strongly inflated to form semicircular lobe. Second maxilliped with basis-ischium fusion incomplete. Third maxilliped with basis-ischium fusion incomplete; basis with one tooth; ischium with crista dentata moderately well developed, with one (occasionally 2) ac- cessory teeth; merus with small spine at dorsodistal margin; carpus un- armed. Sternite of mxps; unarmed, or with spinule on one or both sides of midline; anterior margin with long setae. Right cheliped considerably larger than left; with moderately long, fine setae dorsally. Dactyl equalling, or slightly longer than, length of palm, mesial margin curved; cutting edge with strong calcareous teeth proximally, small calcareous teeth distally; terminating in strong calcareous tooth, slightly overlapped by fixed finger; mesial margin with row of conical spines, decreasing in size distally; dorsal surface with median longitudinal row of conical spines, and with low tubercles toward cutting edge; mesial face between spine rows closely set with tubercles, the more proximal ones produced into small spines; ventral surface with close-set tubercles, and with row of a few spines proximally at ventromesial margin. Palm % length of carpus; lateral margin curved; dorsal surface with irregular rows of high, conical spines, 2 median rows of somewhat larger spines forming inverted V, from its apex a row of enlarged spines extending to tip of fixed finger; dorsomesial margin with row of enlarged spines; dorsolateral mar- gin with row of strong spines, increasing in size on fixed finger; mesial face with close-set, pointed tubercles, these becoming flattened on ventral surface; cutting edge of fixed finger with calcareous teeth, decreasing in size distally. Carpus equalling, or slightly longer than length of merus; dorsal surface convex, with short spines tending to become very small medially; distal margin with row of small spines, dorsomesial margin with row of strong spines increasing in size distally; mesial and lateral faces with close-set tubercles and moderately long setae, distal margins each with row of spines or spinose tubercles; ventral surface tuberculate. Merus sub- VOLUME 90, NUMBER 3 653 triangular; dorsal surface with transverse multidenticulate ridges, distal mar- gin with row of strong spines decreasing in size laterally and mesially; lateral and mesial faces nearly smooth, granulate toward ventral margins; ventro- lateral and ventromesial margins each with row of small spines or spinose tubercles; ventral surface tuberculate. Ischium with row of small denticles or tubercles and tufts of moderately long setae on ventromesial margin. Left cheliped moderately short, sometimes slightly overreaching base of dactyl of right cheliped. Chela subtriangular. Dactyl about 1% times length of palm; cutting edge with row of small corneous teeth; terminating in small corneous claw, overlapped and overreached by fixed finger; dorsomesial margin with double row of strong spines and with tufts of long setae, dorsal surface with flattened granules; mesial face smooth, with long tufts of setae; ventral face smooth or with small pointed tubercles, and with long tufts of setae. Palm slightly more than half length of carpus; dorsal surface flattened, horizontal, with longitudinal rows of conical spines interspersed with small pointed granules and tufts of moderately long setae, one row of spines extending to tip of fixed finger; dorsolateral mar- gin slightly raised, strongly convex, with row of strong spines decreasing in size on fixed finger; dorsomesial margin somewhat expanded, with row of large spines and with tufts of long setae; mesial face with a few tubercles and with tufts of setae; ventral and lateral surfaces with pointed tubercles of various sizes and with tufts of setae; cutting edge of fixed finger with small calcareous teeth, tip with corneous claw. Carpus subtriangular, slightly exceeding merus in length; dorsal surface oblique, with longi- tudinal rows of strong spines and tufts of moderately long setae, distal margin with 2 or 3 rows of small spines, dorsomesial margin with row of strong spines increasing in size distally; mesial and lateral faces with scattered tubercles and tufts of moderately long setae, distal margins each with row of small teeth or tubercles; ventral surface tuberculate. Merus subtriangular; dorsal surface with transverse, multidenticulate ridges and tufts of setae; distal margin with 1 or 2 spines; lateral face nearly smooth, ventrolateral margin with 3 or 4 strong spines distally; mesial face nearly smooth except for several groups of pointed tubercles near proximal end of ventromesial margin; ventromesial margin with row of pointed tubercles and long setae, similar tubercles and setae covering distal part of ventral surface. Ischium with row of strong denticles or pointed tubercles and tufts of long setae on ventromesial margin. Second pereiopods long, frequently reaching or slightly overreaching dis- tal end of right cheliped. Dactyl long, 1*s to 2 times length of propodus; in lateral view, curved ventrally; in dorsal view, twisted; terminating in strong corneous claw; row of calcareous spinules on proximal halt of dorsal surface, this surface with fringe of relatively short setae; mesial face with median longitudinal sulcus, row of corneous spinules close to dorsal margin, 654 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON and a few scattered corneous spinules near ventral margin; lateral surface with median longitudinal sulcus along proximal % to % of segment; ventral margin with row of corneous spines, increasing in size distally. Propodus equalling or slightly exceeding length of carpus; dorsal and ventral margins somewhat converging distally; dorsal surface with row of moderately strong spines; mesial surface nearly smooth, a few low pointed tubercles dorsally and spine on dorsal part of distal margin; lateral surface nearly smooth, a few low tubercles and tufts of short setae dorsally; ventral surface with a few scattered, minute corneous spinules. Carpus % to “4o length of merus; dorsal surface with row of moderately strong spines, increasing in size distally; mesial face nearly smooth, with a few pointed granules near spine row; lateral face nearly smooth, with scattered tufts of short setae and a few rounded granules near spine row; ventral surface smooth. Merus laterally compressed; dorsal margin with row of pointed granules and tufts of moderately short setae, lateral and mesial faces smooth; ventral margin with row of small pointed tubercles, ventrolateral margin with strong spine distally. Ischium with a few small pointed tubercles and tufts of moderately long setae on ventral margin. Third pereiopods approximately as long as second, reaching or slightly overreaching distal end of right cheliped. Dactyl as in Pz. Propodus as in Ps, but spines on dorsal surface smaller and distal margin of mesial surface unarmed. Carpus equalling length of merus; dorsal surface with row of spines, smaller than those of Ps, increasing in size distally; lateral face nearly smooth, with patch of setae dorsodistally; mesial and ventral sur- faces smooth. Merus laterally compressed; dorsal margin nearly smooth, with tufts of moderately short setae; lateral and mesial faces smooth; ventral margin with a few wide-set, small pointed tubercles, ventrolateral margin unarmed distally. Ischium unarmed ventrally. Fourth pereiopods without preungual process on lateral face of dacty]; propodal rasp well developed. Fifth pereiopods typical of Pagurus. Sternite of third pereiopods rounded, anterior margin unarmed but with row of long setae. Pleopods of male unpaired, pl;-pl; with exopodites moderately well developed; endopodites reduced. Pleopods of female unpaired, ple—-plL, with both rami well developed; pl; with exopodite moderately well developed, endopodite reduced. Telson with posterior lobes asymmetrical, left larger than right; sub- quadrate; separated by narrow median cleft; both terminal and both lateral margins with narrow, close-set, strong spines; anterior lobes unarmed, lateral margins with setae. Color in life—Shield and ocular acicles tan, with small, irregularly ar- VOLUME 90, NUMBER 3 655 ranged, reddish-brown flecks. Ocular peduncles white on dorsal and mesial surfaces, reddish-brown on lateral and ventral surfaces; with touches of reddish-brown dorsally at base of peduncle and adjacent to cornea; cornea greenish-black. Antennular peduncles with proximal segments white or pale gray; ultimate segment and flagellum red or reddish-brown. Antennal peduncles with proximal segments white, purplish-brown laterally; ulti- mate segment with sharp line of reddish-brown on lateral and mesial mar- gins; acicle white; flagellum reddish on lateral and tan on mesial side of each article. Chelipeds with ground color tan; spines white at base, purple at tips. Fingers with row of blue tubercles next to cutting edge; to outside of these a narrow, bluish- or purplish-white longitudinal line. Carpus with large, dark purple spot proximally on dorsal surface. Merus with triangular, reddish-brown area dorsodistally; on lateral face, a narrow band of reddish- brown close to distal margin; distal spines reddish-brown with white tips. Ambulatory legs tan. Carpus with large reddish-brown blotch proximally on lateral surface. Merus with broad reddish-brown band at distal end. Etymology—From Greek “spilos,” a spot or stain, and “carpus,” in reference to the distinctive purple mark on the carpal segment of the chelipeds. Remarks.—Available data suggest that this species usually occurs on a sandy substrate. Ovigerous females (11.2-14.9 mm) were collected in April, May, July, and August. Pagurus spilocarpus superficially resembles another Californian species, P. armatus (Dana). In P. armatus, as in the new species, there are spines on the dorsal surface and margins of the chelae; the dorsal surface of the left chela is flattened; the dactyls of the walking legs are twisted, with a ventral row of corneous spines; and there is a dorsal row of spines on the propodus and carpus of the walking legs. However, P. armatus differs in a number of characters, especially in the structure of the ocular peduncles and acicles, left chela, telson, and mouthparts (cf. McLaughlin, 1974: Figs. 13, 14). In the eastern Pacific, Pagurus spilocarpus is most closely related to P. smithi (Benedict). P. smithi appears to be restricted to northern Mexico; its known range (based on published records and a long series of speci- mens in the collections of the Allan Hancock Foundation) includes the en- tire Gulf of California and the outer coast of Baja California as far north as Islas San Benito. This species has not yet been illustrated, but it is easily recognizable from the original description (Benedict, 1892: 4). Among the characters which it shares with P. spilocarpus are the following: The left chela is flat, horizontal, and strongly convex laterally; the telson is asymmetrical, with the posterior lobes subquadrate and armed on both 656 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON | terminal and lateral margins; and the dactyls of the walking legs are twisted, | with a ventral row of corneous spines. There is also a striking similarity in the structure of the mouthparts. These characters are also present in two species from the Atlantic coast of North America, Pagurus pollicaris Say and P. impressus (Benedict) (Wil- liams, 1965: 128, Fig. 103 and 129, Fig. 104). The similarities of these 4 species, and the combined characters which set them apart from other American Pagurus, appear to justify their inclusion in a distinct species- _ group, similar to the 8 groups already recognized and characterized by McLaughlin (1974, 1975) and the one tentatively established by Haig (1974). This may be called the “smithi” group and defined as follows: Anterior margin with rostrum produced, obtusely triangular or broadly rounded. Ocular peduncles stout or moderately stout, corneae dilated. Ocular acicles subtriangular, dorsal surface concave. Chelipeds grossly unequal; left chela with dorsal surface flattened, lat- eral margin strongly convex. Second and third pereiopods elongate; dactyls long, broad, twisted, ventral margins with strong, corneous spines. Fourth pereiopods without preungual process; propodal rasp well developed. Males with 3 unpaired pleopods; females with 4 unpaired pleopods. Telson with posterior lobes asymmetrical, left larger than right; gen- erally subquadrate; terminal and lateral margins with close-set spines. Maxillule with proximal endite subquadrate; endopodite with external lobe produced, ovoid, not recurved. First maxilliped with basal segment of exopodite very strongly inflated, forming semicircular lobe. Second maxilliped with basis-ischium fusion incomplete. Third maxilliped with 1 or 2 accessory teeth on moderately well developed crista dentata; merus with spine at dorsodistal margin. This group has several characters in common with the “bernhardus” group (McLaughlin, 1974: 44), particularly the flattened dorsal surface of the left chela and the form of the rostrum, dactyls of pereiopods 2 and 3, propodus and dactyl of pereiopod 4, and second and third maxillipeds. In members of the “bernhardus” group, on the other hand, the ocular acicles are subovate instead of triangular; the lateral margin of the left chela is not strongly convex; the posterior lobes of the telson are sub- triangular instead of subquadrate, and are unarmed on their lateral margins; on the endopodite of the maxillule, the external lobe is recurved and with- out the distinctive ovoid shape found in the “smithi” group; the basal seg- ment of the exopodite of the first maxilliped is inflated, but much less so than in the “smithi” group and not in the form of a semicircular lobe. Literature Cited Benedict, J. E. 1892. Preliminary descriptions of thirty-seven new species of hermit crabs of the genus Eupagurus in the U.S. National Museum. Proc. U.S. Nat. Miss ola l—26: VOLUME 90, NUMBER 3 657 Haig, J. 1974. Two new species of Pagurus from deep water off Peru and Chile (Decapoda, Anomura, Paguridae). Crustaceana 27:119-130. McLaughlin, P. A. 1974. The hermit crabs (Crustacea Decapoda, Paguridea) of northwestern North America. Zool. Verhandel. Leiden 130:396 pp., 1 pl. 1975. Biological results of the University of Miami deep-sea expeditions. 114. On the identity of Pagurus brevidactylus (Stimpson) (Decapoda: Paguridae), with the description of a new species of Pagurus from the western Atlantic. Bull Mar, Sci. 25:359=376: Williams, A. B. 1965. Marine decapod crustaceans of the Carolinas. U.S. Fish and Wildlife Service, Fishery Bull. 65(1):xi + 298 pp. Allan Hancock Foundation, University of Southern California, Los Angeles, California 90007. PROC. BIOL. SOC. WASH. 90(3), pp. 658-668 ORCHESTIA VAGGALA, A NEW LAND-HOPPER FROM THE GALAPAGOS ISLANDS (CRUSTACEA: AMPHIPODA: TALITRIDAE) Thomas E. Bowman During his recent intensive collecting of land snails throughout the Galapagos Islands, Dr. Joseph Vagvolgyi found a few terrestrial amphipods at three localities at San Cristobal Island. He kindly turned them over to me for study, and I found them to represent a species new to science which is described below. Orchestia vaggala, new species Figs. 1-4 Material examined.—San Cristobal I. (= Chatham I.), Galapagos Is., S Slope of El Junco Peak, near Chino, leg. Joseph Vagvolgyi, Feb. 1974.— Sta. 101, 4 Feb.: 9.4mm ¢ (holotype, USNM 169045); 11.2 mm 2, oostegites with marginal setae; 9.3 mm ovig. 2; 9.3 mm 2, oostegites without setae.— Sta. 103, 4 Feb.: 9.7 mm ovig. 2; 9.5 mm non-ovig. 2.—Sta. 105, 5 and 7 Feb.: 8.1mm 8, oostegites with setae; 8.3 mm 2, oostegites without setae; 9.1 mm 2, oostegites without setae. The 2 listed above are paratypes, USNM 169046 (Sta. 101); USNM 169047 (Sta. 103); USNM 169048 (Sta. 105). Etymology—A combination of contractions of Vagvolgyi, the collector, plus Galdpagos, the locality. Description—Length (head-telson) 8.1-11.1 mm. Eyes black, nearly round, diameter almost 0.4 length of head; separated dorsally by about 0.4 eye diameter. Antenna 1: Not reaching distal margin of antenna 2 peduncle; segments of peduncle subequal in length; flagellum 3-4-seg- mented. Antenna 2: Reaching posterior margin of pereonite 3 to pereonite 5; segment 4 of peduncle about 0.7 length of segment 5; flagellum 14-18- segmented. Mandible: Incisor of left mandible 6-cuspate, of right 5-cuspate; left lacinia with 4 rounded cusps, right lacinia with 3 cusps, more pointed; right spine-row with 6 spines, left with 4; molar with usual transverse rugae on chewing surface, a single guide seta dorsally, and ventrally a flap armed apically with plumose setae. Maxilla 1: As in other species of Orchestia; outer lobe with 9 dentate apical spines; inner lobe with 2 plumose apical setae. Maxilla 2: As in Fig. 1F. Maxilliped: Inner lobe with 3 stout spines and numerous plumose setae on distal margin; palp 3-seg- mented, 3rd segment with minute process apparently representing rudi- mentary 4th segment. VOLUME S0, NUMBER 3 659 Fig. 1. Orchestia vaggala. A, @ head, lateral; B, ¢ right antenna 1; C, ¢ left an- tenna 2, proximal segments; D, Same, distal segments; E, 2 maxilla 1; F, 2 maxilla 2; G, 2 maxilliped; H-I, Same, apex of palp with and without setae; J, Same, apex of inner lobe; K, Pleonal epimera. Fig. 2. A-K, Orchestia vaggala. A-E, @ left mandible: A, Entire mandible; B, Incisor; C—D, Incisor and lacinia, from different aspects; E, Spine-row. F-G, @ right mandible; F, Incisor, lacina, and spine-row; G, Incisor. H, 2 left pereopod 1; I, Same, dactyl and distal part of propus; J, Same, dactyl; K, ¢ pereopod 1. L. Orchestia costaricana, 6 syntype: pereopod 7, basis. Pereopod 1: Nearly identical in ¢ and 2, with only minor differences in setation. Posterior pellucid process of merus small, that of carpus larger; propus widening slightly distally, palm nearly vertical, slightly convex pos- teriorly, overlapped by 2 groups of submarginal spines; dactyl armed as VOLUME 90, NUMBER 3 661 Fig. 3. A-D, Orchestia vaggala. A, 2 pereopod 2; B, Same, dactyl; C, ¢ pereopod 2, lateral; D, Same, medial. E, Orchestia costaricana, 2 syntype: pereopod 2 dactyl. in Fig. 2J. 2 Pereopod 2: Propus widening only slightly distally, extending beyond dactyl by length of latter; palm with 4 submarginal spines overlap- ping dactyl; dactyl thumb-shaped, lateral surface with row of spines in- creasing in length distally. ¢ Pereopod 2: Propus about twice as long as wide, with slightly concave posterior margin; palm transverse, slightly convex, produced into bluntly triangular distal process overlapping tip of dactyl medially, armed with row of short submarginal setae on medial and lateral surfaces; dactyl evenly curved, extending slightly beyond palm, pos- terior margin slightly convex proximally. Pereopods 3 and 4: Slender; dactyl moderately long, nail and proximal part subequal in length. Pereopod 4 of one 2 appeared abnormal, with short carpus and dactyl (Fig. 4C, D). Pereopods 5 and 6: Similar, but pereopod 5 much shorter. Pereopod 7: Posterior expansion of basis with a 662 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON oe Fig. 4. A-P, Orchestia vaggala, 2. A, Pereopod 3, lateral; B, Same, distal end; C, pereopod 4; D, Same, distal end, showing abnormal dactyl; E, Pereopod 5; F, Pereopod 6; G, Same, distal end; H, Pereopod 7; I, Pleopod 1, posterior; J, Pleopod 2, anterior; K, Pleopod 3, anterior; L—N, Uropods 1-3, dorsal; O, Telson, dorsal; P, Same, apex. QO, Orchestia costaricana, 6 syntype: Telson, dorsal. few weak serrations on margin; posteroproximal corner not reaching prox- imal margin of ischium. Marginal setae of oostegites not curl-tipped. Pleonal epimera: Margins unarmed; posteroventral corners angular. VOLUME 90, NUMBER 3 663 Pleopods: Well developed; peduncles with 2 retinaculae; rami 6—10-seg- mented. Uropod 1: Peduncle % longer than rami, with 2 dorsal rows of 3-4 spines and well developed interramal spine; outer ramus with unarmed mar- gins and 4 spines near apex; inner ramus with 4 spines on medial margin and 5 spines near apex, medial apical spine very large. Uropod 2: Peduncle with 4 dorsal spines; each ramus with 2 marginal spines; outer ramus with 4, inner ramus with 5 spines near apex. Uropod 3, peduncle 2.5 times as long and much broader than ramus, with long distolateral spine; ramus with longer lateral spine and minute medial spine on distal margin. Telson suboval, % longer than wide; apex with short rounded notch; dorsal surface with 10 bifid spines arranged as in Fig. 40. Relationships Hurley (1957, 1959, 1968) and others have distinguished between “leaf- mold” talitrids or “land-hoppers,” inhabitants of the forest floor or grass- lands, and supralittoral talitrids or “beach-hoppers” and “rock-hoppers,”’ which live on seashores from low tide level to the spray zone (and some- times above). Following Stebbing’s (1906a) separation of Parorchestia from Orchestia, there was a tendency to assign land-hoppers to Parorchestia and beach-hoppers to Orchestia. The validity of Parorchestia has been some- what insecure, and many authors have treated it as a synonym of Orchestia (e.g. Shoemaker, 1942; Hurley, 1957; 1959; 1968; Barnard, 1958; 1969). In recent years Boustield (1964; 1971; 1976) has recommended that Par- orchestia be recognized as valid and restricted to species of the tenwis-type. He has predicted that new genera will be needed eventually for some land- hoppers now assigned to Orchestia but which differ widely from the type- species, O. gammarellus. The genus Orchestia s.]. (i.e., including Parorchestia), is very large, com- prising more than 100 nominal species. I have not compared O. vaggala with all the known species, but have assumed that the beach-hoppers, which occupy quite a different habitat and must have very different physio- logical requirements, can be all safely excluded from consideration. There remain about 40 species of land-hoppers, now assigned to Orchestia and Parorchestia. The species of Parorchestia can be eliminated because O. vaggala lacks the 4th segment of the maxilliped palp and has normal rather than curl-tipped oostegite setae. Most of the remaining species of Orchestia differ from O. vaggala by one of the following: 1. Lack of an interramal spine on uropod 1 (about 9 species, mostly in the O. floresiana group); 2. Presence of spines on margins of outer ramus of uropod 1 (e.g. kaalensis, luzonensis, notabilis); 3. One or more pleopods reduced to stumps (e.g. lesliensis, patersoni, rubroannulata). The few species still not elim- inated differ in the shape and armature of the telson and or pereopods | and 2 of one or both sexes. 664 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON O. vaggala does not appear to be closely related to any of the known species of Orchestia. A possible candidate for the closest relative is O. | costaricana Stebbing (1960b), known from the mouth of Rio Jesus Maria, Costa Rica, which drains into the Gulf of Nicoya, Puerto Rico (Shoe- maker, 1935), and Santa Cruz I. and Isabela I. in the Galapagos (Monod, 1970). In Costa Rica it was found “in mangroves in the mud under trunks of trees’; in the Galapagos it was collected 1-4 km inland, in soil and crevices. The habitat of the Puerto Rican specimens is unknown. At first glance O. costaricana appears to be far removed from O. vaggala, especially because of its chelate male pereopod 2, but also because of its barely subchelate female pereopod 1 and more sparsely setose maxilliped. On the other hand, there are impressive similarities in other characters traditionally considered to be important in the taxonomy of Orchestia: the male pereopod 1, the basis of pereopod 7 (compare Figs. 2L and 4H), the uropods, and the telson (Fig. 40, Q). Further speculation on the relationships of O. vaggala will not be at- tempted, since many land-hopper species on Pacific Islands await discovery (Bousfield, 1964; 1971; 1976) and since endemic, undescribed land-hoppers occur on Jamaica, Haiti, and Barro Colorado Island, Panama (Hurley, 1968). Habitat San Cristobal is the easternmost and the fifth largest of the Galapagos Islands, having a surface area of 558 km? and a maximum elevation of 730 m. A prominent feature is the crater on the southern part of the island that contains a permanent freshwater lake, El Junco. This lake lies at an eleva- tion of about 700 m and covers most of the floor of the crater (maximum diameter 290 m) with 6 m of water, a depth established by a low point in the crater rim where the lake overflows (Colinvaux, 1968). Water from El Junco and probably from underground sources also feeds intermittent streams on the south slope of the peak, as well as the only permanent stream on the archipelago, which empties into Bahia de Aquadulce (Freshwater Bay) on the southern coast. Precipitation varies greatly from year to year (Fig. 5), but in general there is a wetter season from December to May and a drier season from June to December. During the latter, a fine mist covers the heights, and although it does not fall as rain, it thoroughly wets the vegetation (Thornton, 1971). It is evident, therefore, that the southern slope of El Junco peak provides enough soil moisture for the needs of land-hoppers. It is probably not just by chance that although Dr. Vagvolgyi collected land snails at many localities throughout the Galapagos Islands, he encountered land-hoppers only on San Cristobal. Collection data for the 3 sites where Orchestia vaggala was found is as follows: VOLUME 90, NUMBER 3 665 Fig. 5. Precipitation (mm) at El Progreso, San Cristébal I. (west of El Junco peak), 1966-1970. Redrawn from Comision Interinstitucional etc. (1973). Sta. 101. Elevation 555 m. Miconia-zone. Collections made on bottom and sides of a wash, both from the litter and from arboreal mosspads; amphipods found only on the ground. In the wash was a dense growth of Miconia robinsoniana bearing many epiphytes: mosses, lichens, liverworts, and ferns. Ferns were also growing on the ground, and there was an occasional guava tree (Pisidium guajava). The ground was covered with brown soil with an occasional rock. Sta. 103. Elevation less than 450 m. Probably in Miconia-zone, but possibly in moist forest zone. In same wash as Sta. 101 but farther down- stream. Sta. 105. Elevation between 550 and 600 m. Fern-sedge zone. Collec- tions made at base of clumps of ferns (Pteridium aquilinum), grasses, and sedges. Trees absent but tree-fern Cyathea weatherbyana was grow- ing in washes or steep areas. The ground was covered with soil; rocks were present only in the washes or on the steepest walls. Origin Five possible explanations for the occurrence of O. vaggala on San Cristobal I. merit discussion. 1. O. vaggala developed locally from an unknown beach-hopper. Dr. J. L. Barnard kindly allowed me to search his Galapagos collections for talitrids. None of the beach-hoppers that I found were species of Orchestia, hence a Galapagos beach-hopper similar to O. vaggala, if it exists, remains to be found. Bousfield (1968) suggested that the less specialized land hoppers (which would include O. vaggala) evolved directly from seashore 666 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON : a. JUNCOO) acy - ONS 5@ °° ACHIN f=" EL PROGRESOA We @ 103 Fig. 6. San Cristdbal I., showing collection stations (solid circles) and other lo- calities mentioned in text. Arrow points to Bahia de Aquadulce. Scale = 5 nautical miles. species, whereas the more modified species “most likely originated in contiguous continental land masses centering on Antarctica, probably sometime during the Mesozoic... .” 2. O. vaggala was brought to San Cristdbal I. on vegetation introduced by man. 3. O. vaggala was carried to San Cristobal I. on a raft of drifting vegeta- tion. Thornton (1971) discusses the possibility of such rafts reaching the Galapagos from Peru and Ecuador with the Humboldt Current or from Panama with the South Equatorial Current. 4. O. vaggala was transported to San Cristébal by birds. Observations and experiments by Segerstrale (1954) support the possibility of dispersal on ducks of Gammarus lacustris, but not G. pulex. Transport of Hyalella azteca by mallard ducks is documented by Rosine (1960). The possibility that O. vaggala was carried to San Cristébal I. by birds seems unlikely, con- sidering the distance involved, but cannot be dismissed outright. Some support for the possibility is given by the fact that numerous waterfowl, including ducks and curlews, live in El Junco. d. O. vaggala migrated from western South America to the coastal “an- cestral Galapagos Is.” and dispersed gradually along island stepping stones VOLUME 90, NUMBER 3 667 as easterly components of the archipelago subsided and more westerly islands emerged (Holden and Dietz, 1972; Rosen, 1975). This view of the history of the Galapagos contrasts with the view that these islands, about 1 million years old, are truly oceanic and have never been con- nected to or in proximity to a continental land mass. It also allows much more time, up to 40 million years, for the evolution of endemic species on the ancestral and modern Galapagos Is. Acknowledgments I am most grateful to Dr. Joseph Vagvolgyi for donating his specimens to the Smithsonian Institution and allowing me to study them, and for reviewing the manuscript. Dr. Thomas E. Simkin generously provided me with literature and information on the geography of San Cristobal. Dr. J. L. Barnard offered helpful advice and reviewed the manuscript. Literature Cited Barnard, J. Laurens. 1958. Index to the families, genera, and species of the gam- maridean Amphipoda (Crustacea). Allan Hancock Foundation Publications, Oc- casional Paper No. 19:145 pp. —. 1969. The families and genera of marine gammaridean Amphipoda. Bulletin of the United States National Museum 271:vi + 535 pp. Bousfield, E. L. 1964. Insects of Campbell Island. Talitrid amphipod crustaceans. Pacific Insects Monograph 7:45-57. 1968. In: Session I: Discussion, Transition to land. American Zoologist 8(3):393-398. —. 1971. Amphipoda of the Bismarck Archipelago and adjacent Indo-Pacific islands (Crustacea). Steenstrupia 1:255-293. —. 1976. A new terrestrial amphipod from Lord Howe Island. Records of the Australian Museum 30(6):118—122. Colinvaux, Paul A. 1968. Reconnaissance and chemistry of the lakes and bogs of the Galapagos Islands. Nature 219(5154):590-594. Comsién Inter-institucional Enviada por el Gobierno del Ecuador al Archipiélago de Colon, desde el 20 de Enero al 16 de Febrero de 1973. 1973. Recursos del Archipiélago de Colén, estado actual de aprovechamiento y posibilidades de desarrollo. Quito, Ecuador, viii +- 146 pp, 34 anexos, 3 graphs, 2 maps [mimeo- graphed]. Holden, John C., and Robert S. Dietz. 1972. Galapagos gore, NazCoPac triple junction and Camegie/Cocos Ridges. Nature 235(5336):266—-269. Hurley, Desmond E. 1957. Terrestrial and littoral amphipods of the genus Orchestia, family Talitridae. Transactions of the Royal Society of New Zealand 85(1): 149-199. 1959. Notes on the ecology and environmental adaptations of the terres- trial Amphipoda. Pacific Science 13(2): 107-129. 1968. Transition from water to land in amphipod crustaceans. American Zoologist 8(3):327—353. Monod, Théodore. 1970. Sur quelques Crustacés Malacostracés des Tles Galapagos récolté par N. et J. Leleup. Mission Zoologique Belge aux Tles Galapagos et en Ecuador (N. et J. Leleup, 1964-1965). 2:11-53. 668 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Rosen, Don E. 1975. ) Ske 03o5— N=4 N=2 0.880 0.500 0.885 0.406 Ng 10 Ne 0 Nij=10 N = 10 Rana 43.9— 50.7 (Tympanum M = (0.458 M = 0.971 M0327 kuhli 74.3 67.4 hidden) R =0422— RR: = 0905— Re 103307— ING p13 Nia 0.481 1.09 0.351 N15 Nes NaS * A female at 67 mm is immature. erately or deeply pigmented with slate-brown blotches; throat and head region usually dark slate-brown. Measurements of holotype (mm).—Snout-vent length 63.1; length of head 23.5; breadth of head 24.8; length of snout 8.8; diameter of eye 120: diameter of tympanum 3.0; interorbital breadth 4.6; length of third finger 9.8; length of second finger 7.0; length of first finger 8.1; diameter of fourth toe disk 2.5; length of tibia 30.3; length of hind limb 93.5. Ecological note —One specimen was collected in the water and the others on rocks in the Tagibo River, a gradient stream, in an area of dipterocarp forest, at an altitude of about 1,000 meters. Range.—Known only from the type locality in northeastern Mindanao. Although we have conducted extensive surveys on western Mindanao and the nearby islands of Camiguin, Bohol, Cebu, Siquijor and Negros, this species has not been taken in these localities. Its possible occurrence on Dinagat, Samar, or Leyte remains to be determined. Etymology.—The name “diuata’ is that of the mountain range in eastern Mindanao where the frog was discovered. Comparison with other species.—Rana diuata is distinguished from Rana magna, the sympatric population with which it might be confused, by its smaller size; darker, more uniform dorsal coloration; more rugose skin on the anterior dorsal surfaces; shorter first finger, about equal to the sec- VOLUME 90, NUMBER 3 673 Fig. 2. Rana magna: a, Palmar view of hand; b, Plantar view of foot. ond when fingers are adpressed (not distinctly longer than); somewhat more dilated toe disks (Figs. 1 and 2) and shorter tibia relative to the snout- vent length (see Table 1). From Rana woodworthi which occurs on Luzon and Polillo Islands, it can be distinguished by its very much darker and much more uniform coloration; slightly less expanded finger tips; and much more tuberculate skin (R. woodworthi is relatively smooth); shorter first finger, about equal to the second when adpressed (not longer than); and the ab- sence of a dorsolateral fold which is prominent for R. woodworthi. This species in habitus, size, and general dorsal color pattern, webbing, tubercu- lation and habitat preference is most like R. kuhli from Borneo and southeast Asia. It differs in a number of characteristics: the distinct (not hidden) tympanum (Fig. 3), the longer snout as evidenced by the ratio of snout length to head length and the diameter of eye to snout length (Table 1), the more pointed snout when viewed dorsally, and the darker ventral color. 674 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 3. Head of: a, Rana diuata; b, Rana kuhli. Detailed descriptions of R. magna, R. woodworthi and R. kuhli are avail- able in recent publications (Inger, 1954, 1958, and 1966). Acknowledgments We are indebted to Robert F. Inger and Hyman Marx for assistance in the comparison of this species with R. kuhli from Borneo. Illustrations were prepared by Walter Zawojski, Stanford University. This study is part of our Philippine-Pacific Islands research project under the auspices of National Science Foundation Grant GB-41947. VOLUME 90, NUMBER 3 675 Literature Cited Alcala, Angel C. 1962. Breeding behavior and early development of frogs of Negros, Philippine Islands. Copeia 1962:679-726. Inger, Robert F. 1954. Systematics and zoogeography of Philippine amphibia. Fieldiana, Zool. 34:183-531, Fig. 39-98. ——., 1958. A note on Philippine frogs related to Rana macrodon. Fieldiana, Zool. 39:253—-255. ——. 1966. The systematics and zoogeography of the amphibia of Borneo. Field- Tana ZOO 521-402, Eos (ln (WCB) Menlo College, Menlo Park, California and Department of Herpe- tology, California Academy of Sciences, San Francisco, California; (ACA) Department of Biology, Silliman University, Philippines. PROC. BIOL. SOC. WASH. 90(3), pp. 676-684 SARSIELLA MAURAE, A NEW SPECIES OF MARINE OSTRACODA (SARSIELLIDAE: MYODOCOPINA) FROM BAHIA DE LOS ANGELES, GULF OF CALIFORNIA, MEXICO Louis S. Kornicker Abstract—Kornicker, Louis §., Department of Invertebrate Zoology, Smithsonian Institution, Washington, D.C. 20560.—Sarsiella maurae, a new species of marine Ostracoda (Sarsiellidae: Myodocopina) is described from Bahia de Los Angeles, Gulf of California, Mexico. This is the first record of the family from the Gulf. This paper describes and illustrates a new species of benthic myodocopid ostracode from Bahia de Los Angeles, Gulf of California, Mexico. No myodocopid Ostracoda have been described previously from the Gulf of California. Bahia de Los Angeles is a small bay along the eastern shore of Baja California. Barnard and Grady (1968) have presented a general account of the bay environment. Shore sediments consist mostly of shell fragments and rocks. The available data for 1962, the year in which the species de- scribed herein was collected, show a low temperature of less than 15°C in February and a high of 29.8°C in August. Salinities in the bay in April were recorded as 35.25% at the surface and 35.10% at a depth of 30 m. Suborder MYODOCOPINA Sarsiellidae Brady and Norman, 1896 Sarsiella Norman, 1869 Sarsiella maurae, new species Figs. 1-6 Etymology.—The species is named for Maura McManus, Smithsonian Institution, who assisted in the preparation of the manuscript. Holotype —USNM 156739, unique specimen, adult or A-1 female, length 1.22 mm, on slides and in alcohol. Type-locality—Bahia de Los Angeles, station SIO-62-216, 21 April 1962, shore sample, reef between Isla Ventana and Isla Cabeza de Caballo, tailings of fish sample. Collected by Dr. Carl L. Hubbs and party, Scripps Institu- tion of Oceanography. Specimen received from Dr. J. Laurens Barnard. Description of female (either adult or A-1 instar)—Carapace oval in lat- eral view with small posteroventral caudal process (Figs. 1, 2); carapace broadest in posterodorsal part (Fig. 3a). Ornamentation (Figs. 1-4b, c): Each valve with 7 ribs radiating from hub in area of central adductor muscle attachment (Figs. 1, 2); rib extend- ing onto posterodorsal part of valve terminating at low ridge and more VOLUME 90, NUMBER 3 67 “I Fig. 1. Sarsiella maurae: Lateral view of complete specimen, length 1.22 mm. prominent than other ribs; ventral, anterior, and anterodorsal ribs each terminating in small marginal process; 1 or 2 small marginal processes present between rib terminals; posterior of valve dorsal to caudal process with 4 small marginal processes; an additional small process present on posterodorsal margin; several shallow fossae present at hub of radial ribs; minute spines present on caudal processes and on marginal processes (Figs. 3d—-g); surface between ribs with minute pits and pebbly texture (Fig. Ac); long bristles along anterior, ventral, and posterior margins of each valve and sparsely distributed on valve surface (Figs, 2, 3h, i, 4b). Infold (Figs. 4a, d-h): Minute bristle near middle of anterior infold; caudal processes with 4 or 5 bristles with single or double pointed tips (Figs. 4e, g, 5a); inner edge of infold of ventral margin near caudal process with 4 or 5 slender bristles with open ends (Figs. 4h, 5a); posterior infold dorsal to caudal process with 2 setose bristles (Figs. 4f, 5a). Size: Holotype, length 1.22 mm, height 1.06 mm. First antenna (Fig. 5b): Ist joint bare; 2nd joint with 1 dorsal bristle and spines along dorsal margin; 3rd and 4th joints fused; 3rd joint with 1 dorsal bristle, but without ventral bristle; 4th joint with spines forming rows on medial surface and along dorsal and ventral margins, and with 3 bristles, 1 dorsal, 2 ventral; 5th joint with spines forming row on distal dor- sal corner; sensory bristle of 5th joint with 1 minute proximal filament and 678 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 2. Sarsiella maurae: Stereoscopic pair showing lateral view of carapace, dorsal to right; length of specimen 1.22 mm. spine at tip; medial bristle of small 6th joint short. Seventh joint: a- bristle about 3 times length of bristle of 6th joint; b-bristle about twice length of a-bristle, with minute spine at tip; c-bristle about same length as sensory bristle of 5th joint, with 1 minute proximal filament and minute spine at tip. Eighth joint; d- and e-bristles bare, slightly shorter than c-bristle; f-bristle shorter than c-bristle, with 1 minute proximal filament and 1 minute terminal spine; g-bristle same length as c-bristle, with 1 minute proximal filament and spine at tip. Second antenna: Protopodite bare. Endopodite 1-jointed, with small terminal protuberance, 2 short anterior proximal bristles, and 1 or 2 short terminal bristles (Fig. 5c). Exopodite: Ist joint with small recurved medial bristle on distal margin; bristle of 2nd joint with about 28 proximal ventral spines and distal natatory hairs; bristles of joints 3-8 with proximal ventral spines and distal natatory hairs; 9th joint with 1 long bristle with few prox- > Fig. 3. Sarsiella maurae: Outside views of right valve: a, Dorsal view, <68; b, An- terior view, 68; c, Posterior view, x68; d, Posterior end of posterior rib, from Fig. 2, 625; e, Process on posterior rib, from d, <7,500; f, Caudal process, from Fig. 2, x360; g, Processes and bristles on caudal process, from f, 3,500; h, Bristle from near middle of carapace, 1,450; i, Base of bristle shown in h, X5,500. Given magnifications are those at which the micrographs were made on SEM; these have been reduced 58% for pub- lication. "on py ibittip \N WN NSS Ga) pC ea) 6a a > Z S o>) = a > =| © = 680 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON imal ventral spines and many distal natatory hairs, and 1 short dorsal bristle with short marginal hairs; basal spines absent; no short spines observed along distal margins of joints. Mandible (Fig. 5d): Coxale endite represented by stout spine; ventral margin of coxale hirsute. Basale: dorsal margin with 1 short bristle at middle and 2 subterminal; medial side with 2 short bristles near ventral margin; ventral margin with 1 minute and | short bristle; lateral surface with 2 short bristles near ventral margin. Exopodite absent. Endopodite: lst joint with spines on medial surface, shorter spines forming medial row on distal margin near dorsal edge, and stout ventral claw; 2nd joint with small, terminal, dorsal bristle and stout ventral claw; 3rd joint with stout claw and 2 minute bristles, 1 ventral and 1 dorsal to base of claw. Maxilla: Protopodite with hairs along dorsal margin; 3 endites with total of about 11 bristles. Basale with 1 short bristle at distal dorsal corner and 1 near base of exopodite. Exopodite with 3 bristles: 1 long and bare, 1 about half length of long bristle and with few spines, and 1 about *% length of short bristle and bare. Endopodite: Ist joint with stout spinous alpha- and beta-bristles and spines along anterior margin proximal to alpha-bristle; 2nd joint with 2 short a-bristles, 1 short c-bristle, and 5 pectinate end bristles. Fifth limb (Fig. 6a): Single endite with 1 short bristle; 2nd to 5th joints fused; 2nd joint with 3 terminal bristles; 3rd to 5th joints with total of 5 bristles; 2nd to 5th joints hirsute. Sixth limb (Fig. 6b): Single endite with 1 long terminal bristle and 2 short proximal bristles; end joint with 13 bristles with faint marginal hairs followed by space and then 2 stout hirsute posterior bristles; limb hirsute. => Fig. 4. Sarsiella maurae: a, Base of upper bristle shown in f, x9,000; b, Base of bristle from near middle of outside of valve, x5,500; c, Detail of surface near middle of outside of valve, 10,000; d, Inside view of valve (top tilted about 30° forward), <65; e, Inside view of caudal process shown in d, x 260; f, Two setose bristles on posterior infold dorsal to caudal process, 1,500; g, Bristle on infold of caudal process shown in e, 3,200; h, Tubular bristles on inner edge of ventral infold just anterior to caudal process, 2,900. Given magnifications are those at which the micrographs were made on SEM; these have been reduced 58% for publication. Fig. 5. Sarsiella maurae: a, Posterior of right valve, inside view; b, Ist antenna, medial view; c, Endopodite of right 2nd antenna, medial view; d, Left mandible, medial view; e, Left lamella of furca, lateral view; f, Anterior of right lamella of furca showing proximal parts of claws 1 and 2, medial view. Fig. 6. Sarsiella maurae: a, 5th limb; b, 6th limb; c, 7th limb; d, Right lateral eye, medial eye and rod-shaped organ; e, Anterior of body showing anterior process and upper lip; f, Left Y-sclerite, lateral view, anterior to left. a) aa ica jaa) = =) Z >) VOLUME 96 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 682 Wy = — - = — = —s — qq itt VOLUME 90, NUMBER 3 684 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Seventh limb (Fig. 6c): Distal group with 6 bristles, 3 on each side; prox- imal group with 3 or 4 bristles, 1 or 2 on each side; each bristle with 3 to 6 bells. Terminus consisting of opposing combs, each with 5 or 6 curved teeth. Furca (Fig. 5e, f): Each lamella with 5 claws followed by 3 or 4 spines; claw 1 united to lamella, claws 2-5 separated from lamella by suture; claws 1-4 with teeth along posterior margins, some longer than others; claw 5 with few teeth along posterior margin; right lamella with few spines along anterior margin proximal to claw 1, and long medial hairs at base of claw 1. Eyes and rod-shaped organ (Fig. 6d): Medial eye with dark brown pigment, bare; lateral eye about same size as medial eye, with 5 ommatidia and dark brown pigment between ommatidia; rod-shaped organ elongate, l-jointed with rounded tip. Upper lip: Helmet shaped with small anterior projection (Fig. 6e). Genitalia: Not observed. Brushlike organ: Not observed. Y-Sclerite: Typical for genus (Fig. 6f). Hees None: Comparisons.—The ornamentation of the carapace of the new species S. maurae differs from that of previously described species of Sarsiella. Also, the 3rd joint of the Ist antenna of S. maurae is without a ventral bristle, whereas previously described species in the family have a ventral bristle on the 3rd joint. However, because only 1 specimen of S. maurae is on hand, it is not possible to ascertain the variability of this character. Acknowledgments I wish to thank the following individuals: Dr. J. Laurens Bamard for submitting the specimen described herein, Carolyn Bartlett Gast for render- ing the illustration of the carapace in Fig. 1, Mr. Walter R. Brown for assisting in the operation of the scanning electron microscope, and Anne C. Cohen for criticizing the manuscript. Literature Cited Barnard, J. Laurens, and John R. Grady. 1968. A biological survey of Bahia de Los Angeles, Gulf of California, Mexico. I. General account. Transactions of the San Diego Society of Natural History 15(6):51—66. Brady, G. S., and A. M. Norman. 1896. A monograph of the marine and freshwater Ostracoda of the North Atlantic and of North-Western Europe. Scientific Transactions of the Royal Dublin Society 5(2):621-684. Norman, A. M. 1869. Shetland Final Dredging Report, Part II: On the Crustacea, Tunicata, Polyzoa, Echinodermata, Actinozoa, Hydrozoa, and Porifera. Pages 247-336 in Report of the Thirty-Eighth Meeting of the British Association for the Advancement of Science. PROG, BIO, SOC, WASH: 90(3), pp. 685-689 PAXONOMIC STATUS OF THE TURTLE, CHRYSEMYS PICTA, IN THE NORTHERN PENINSULA OF MICHIGAN Carl H. Ernst and James A. Fowler The proper subspecific designation of the painted turtles, Chrysemys picta, populating Michigan’s northern peninsula has long been questioned. Conant (1975) showed two subspecies, C. p. bellii and C. p. marginata, occurring there and forming an intergrade population. This was probably based on Ernst (1971) who theorized this area to represent a zone of in- tergradation between these two subspecies. However, until now no study of these turtles from northern Michigan has been reported. Methods and Materials Painted turtles from 12 countries in the northern peninsula of Michigan were examined (Table 1, Fig. 1). Specimens were borrowed from mu- seums, or collected, examined, photographed and released alive by the junior author. Chrysemys p. bellii has alternating vertebral and pleural seams; a re- ticulate pattern of yellow or red lines on the carapace; a large, dark plastral figure, which branches out along the seams and occupies most of the plastral surface; and narrow seam borders on the carapacial scutes which are usually yellow or red (in approximately 80% of specimens examined). Chrysemys p. marginata has alternating vertebral and pleural seams; no reticulate pattern on the carapace; a dark plastral figure which is usually no more than half the width of the plastron and does not extend along the scute seams; and usually (approximately 75% of specimens examined) nar- row olive seam borders on the carapacial scutes. In order to eliminate variation owing to intergradation, we based our subspecific distinctions on 71 C. p. bellii from Murray County, Minnesota (Ernst and Ernst, 1972) and 252 C. p. marginata from Tennessee, Kentucky and Pennsylvania (Ernst, 1970; Ernst and Ernst, 1971). These samples are included in Table 1. The degree of disalignment of the carapacial seams was determined as follows. When the seams between the vertebral and pleural scutes lie in the same transverse line, they are considered 0% disaligned; if the seams alternate exactly they are 100% disaligned. The base point for measuring is the inner end of the seam between pleurals 2 and 3. The imaginary line from the base point forward and parallel to pleurals 1 and 2 is measured and denoted la on the left side and 2a on the right. The part of this same imaginary line starting at the base point and extending forward to the point 686 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 1. Localities of Chrysemys picta examined. See Table 1 for locality names and number of specimens. Solid circles = Chrysemys p. bellii populations; hollow circles = Chrysemys p. marginata populations; and stars = intergrade C. p. bellii x p. mar- ginata populations. opposite the end of the seam between vertbrals 2 and 3 is measured and denoted lb in the left side and 2b on the right. The average percent disalignment is then calculated as 1b/la + 2b/2a (Hartman, 1958). The border of the anterior seam of pleural 2 was measured at its widest point and its color noted. All measurements were taken with dial calipers accurate to 0.1 mm. Results and Discussion Mensural and meristic data of the characteristics separating C. p. bellii and C. p. marginata are given by counties in Table 1 along with similar data from pure populations of these subspecies. The percentage of disalignment of the carapacial seams indicated that the populations, except Ontonagon County (2), more closely resembled C. p. bellii, although each included turtles with disalignment within the range of C. p. marginata. The plastral patterns showed much variation. The number of turtles with the bellii-type pattern decreased from west to east, while those with intergrade or marginata-type patterns increased. The occurrence of a reticulate carapacial pattern, characteristic of C. p. bellii, decreased from west to east. The color of the carapacial seam bor- ders and width of the second pleural seam were variable throughout the region. 687 VOLUME 90, NUMBER 3 OOO CoO) LI CaS GG 0 6S SS Se 7 LOI-8'S3 (0'8) 9°G8 GS Ceg “Ay “uuay ) DIUIsIDU “A “4D e¢-60 (60) 6T OG cL 9°86 > = — OO! L'801-0'6L (18) PLAS We ( BJOSOUUTLYA ) 9g ‘ad’d S90 (Ol) Ee 0G 0&6 O09 0 O83 Of = SP6EO06L (6S) GLB O1 ‘0—D emoddiyy —- “ZT Sol (SO) ei Si == 261 (0) LO SC Ce a 0'06-0'3L (86) $98 9 0D) oeubpeyy Hil Fi-80 OO) VA == as 2 LY) 0 Ole. == F66-09L (TL) S68 € OD 2] ‘OT Per Co) Se Ol OG OL OT OS OV OI 8'66-6'69 (L'8) 0'68 OT “OD Ferojooyos ‘6 II-6l (EO) ot = 008 es G/F) ae we O'36-8'38 (LY) T88 6E ‘0D 10518 SES (eh) Te = NE) ASS 0 oe A) ae 0'00I-0'6L (EIT) BLS € ‘O) ~=9VeIUTULOUI Vl cL Stoo O) (ST) OG See ee O0F 0 OS - QS = O'F6-G'3S (18) G88 G ‘O*—) UOSUPPIC 9g S680 (CGO) Wi 96 96 LP SE VG Gy IE OOII-FIL (931) G6I6 Is ‘0-) sponbieyy g REPT Res = oe OOR 08 —= =e 0G 8'96-G'Sg (FG) S IG ¢ ‘O-) MeuooMoy 4 =e Se 9G = == OOF = 200K = == 1001 =e ss GSS I ‘OD uoWYsnoHH 'E T€-60 (g'0) 6T JEL OS ASS OOT pe a U0 SE6-66S (FSI) LES 9 ‘0F) UosPU0UC G 3 ae VE OS sae OS OOT ae 209° 0S 9°€6-G'gg8 (9S) LauG G ‘OF) 91Ga804) =I esuey (qQ'sSz) ¥ On A uioned wt q apuoy (CPS se) % N Ay[es0'T ‘ON [etoedeieo dey uvas [ened IO[OO UIeaS oem utoyed JUOUIUST[esIp pug wpm jeineid % -o1jo 2%, [eryseld % ulvas [eIoedeies % ‘(QATTO = 0 ‘pal = 1 ‘MOTJPA = A ‘apeisieJUl DyDUIsZIDW X MJaq = 1 ‘DUsIW = W Wag = q) ‘“WUI UI SyUEWAINSeeU [TY ‘“peIpNys sonstiojovreyo OF soiyypeoo, Aq eyep [eosHeIg “T 2qe], 688 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON | The only populations that could be assigned to C. p. bellii were those of Ontonagon and Houghton counties (localities 2 and 3) and, since few specimens were examined from each, they may later prove to include intergrades. Similarly only localities 10 (Luce County) and 12 (Chippewa County) appeared to be C. p. marginata. All other counties (4-9, 11) con- tained intergrade populations (Fig. 1). The study area contains three major drainage systems: Lake Superior; Lake Michigan; Lake Huron. The Lake Superior drainage contained localities, 2, 3, 4, 8, 10 and parts of 5 and 12 (specimens examined from Maraquette County (5) and Chippewa County (12) were from several localities in each and were lumped for reporting ease). Painted turtles in the western parts of this drainage were C. p. bellii, intergrades appeared in the central parts, and, in the east, specimens appeared to be C. p. marginata. The Lake Michigan drainage contained localities 1, 6, 7, 9, 11, and part of 5. All specimens from this drainage were intergrades. The Lake Huron drainage covered only part of locality 12 and these turtles were C. p. marginata. Probably a multiple invasion of the northern peninsula occurred after the final retreat of the Pleistocene glaciers made habitats once more suitable. Chrysemys p. bellii invaded the area from the west (northern Wisconsin and Minnesota); while C. p. marginata probably entered by three routes; the east coast of Wisconsin, the lower peninsula of Michigan by island hopping across the Straits of Mackinac, and from Ontario in the east. Later gene exchange between the two subspecies formed the present zone of inter- gradation in the center of the northern peninsula. Acknowledgments Appreciation is expressed to curators of lending institutions for prompt handling of requested loans and other courtesies. Specimens were borrowed from the following institutions: Chicago Academy of Sciences; Field Museum of Natural History; The Museum, Michigan State University; Museum of Zoology, University of Michigan; and Northern Michigan Uni- versity. Special thanks is given to Dr. J. Kirwin Werner of Northern Michigan University for supplying additional data. Literature Cited Conant, R. 1975. A field guide to reptiles and amphibians of eastern and central North America. Houghton Mifflin Co., Boston. 429 p. Ermst, C. H. 1970. The status of the painted turtle, Chrysemys picta, in Tennessee and Kentucky. Jour. Herpetol. 4:39—45. ——. 1971. Chrysemys picta. Catalog. Amer. Amphib. Rept. 106:1—4. , and E. M. Emst. 1971. The taxonomic status and zoogeography of the painted turtle, Chrysemys picta, in Pennsylvania. Herpetologica 27:390-396. , and 1972. Biology of Chrysemys picta bellii in southwestern Min- nesota. Jour. Minnesota Acad. Sci. 38:77-80. VOLUME 90, NUMBER 3 689 Hartman, W. L. 1958. Intergradation between two subspecies of painted turtle, genus Chrysemys. Copeia 1958:261—265. (CHE) Department of Biology, George Mason University, Fairfax, Vir- ginia 22030; and (JAF) Cranbrook Institute of Science, Bloomfield Hills, Michigan 45013. PROC. BIOL. SOC. WASH. 90(3), pp. 690-697 A GREAT AUK, PINGUINIS, FROM THE PLIOCENE OF NORTH CAROLINA (AVES: ALCIDAE) Storrs L. Olson The Great Auk, Pinguinis impennis, is renowned chiefly for the sad history of its extinction, the last known individuals having been killed in 1844 | (Greenway, 1958). Of the Recent species of Alcidae, P. impennis was the largest and the only one that was flightless. In historic times it is known to have bred on certain North Atlantic islands off Newfoundland, Iceland, and Britain, and to have wintered south to New England in the western Atlantic and to Spain in the eastern Atlantic. Remains indicating a more extensive range in the past have been found in middens in Florida and in the Pleistocene of Italy, as well as at additional localities between these sites and the known historic range (Brodkorb, 1967). Although a number of authors have considered the Great Auk to be con- generic with the modern Razorbill, Alca torda, I agree with Salomonsen (1944) and others that impennis should be separated under the genus Pinguinis. The great modifications seen in the wing of Pinguinis are not the result of neoteny, as seen in many other flightless birds (Olson, 1973), or of “degeneration,” as suggested by Greenway (1958:271), who was under the impression that the keel of the sternum of Pinguinis was as reduced as the wing. Instead, these modifications represent highly derived specializa- tions for wing-propelled diving. Until now there has been no fossil record of Pinguinis other than the Pleistocene occurrences of P. impennis. This lack has been remedied by the discovery, among the thousands of fossil alcid bones recovered from a phosphate mine in North Carolina (Olson and Wetmore, in press), of six specimens that pertain to an undescribed species recognizable as be- longing to the genus Pinguinis by its short, heavy ulna. For comparison I have had the extremely large series of disassociated bones of P. impennis collected by Lucas (1890) on Funk Island, Newfoundland. Pinguinis alfrednewtoni, new species Figs. 1-3 Holotype.—Right ulna lacking only the most distal portions of the condyles, National Museum of Natural History vertebrate paleontological collections, USNM 193334. Locality—Spoil piles at the Texasgulf phosphate mine at Lee Creek, south side of Pamlico River, near Aurora, Beaufort County, North Carolina (35°23’N; 76°47'30’W); collected in October 1973 by Roger C. Wood, et al. Horizon.—Lower part of Yorktown Formation, Chesapeake Group, Lower VOLUME 90, NUMBER 3 691 Pliocene. Strata of both Middle Miocene (Pungo River Formation) and Lower Pliocene (Yorktown Formation) age contribute fossils to the spoil at the Lee Creek mine. Most of the vertebrates are from the Yorktown For- mation and the preservation of the types of P. alfrednewtoni is similar to that of other specimens from this level. Matrix samples from USNM 193334 and 179226 were analyzed by Dr. Thomas G. Gibson, who, on the basis of the foraminiferan fauna and sedimentary characteristics, assigned them to the lower part of the Yorktown Formation. The consensus de- veloped elsewhere is that the Yorktown Formation is Lower Pliocene in age (Ray, in press). Measurements of holotype.—Total length as preserved 54.1 mm (the complete length would probably not have been more than a millimeter greater), depth of proximal end 14.9, width of proximal end 5.1, greatest diameter of internal cotyla, 7.0, width of shaft at midpoint 5.3, depth of shaft at midpoint 8.3, width of distal end 6.9, depth of distal end 10.3. Paratypes.—Distal half of left humerus USNM 179226; very worn distal third of right humerus USNM 192497; left femur abraded at the ends USNM 206362; worn right tarsometatarsus lacking the trochleae USNM 179277; very worn distal fourth of right tibiotarsus USNM 193101. All of these specimens were collected at the same locality as the holotype on various visits by USNM staff or collaborators. Measurements of paratypes—Humerus 179226; greatest diameter of distal end 16.7 mm, depth through internal condyle 12.7, distance from prox- imal end of ectepicondylar prominence to distal end of internal condyle 15.9, least width of shaft 10.3, least depth of shaft 5.4. Femur 206362; overall length 71 mm, width of proximal end 14, width of shaft at midpoint 6.4, depth of shaft at midpoint 7.4. Tibiotarsus 193101; width of distal end c. 12.5, depth through internal condyle c. 11.6. Tarsometatarsus 179277; width of proximal end 13.3, depth of proximal end 11.8, distance from proximal margin of intercotylar prominence to distal foramen 38.9, least width of shaft 6.5, least depth of shaft 5.2. Etymology.—This species is dedicated to the memory of Alfred Newton (1829-1907). Newton was one of the principal founders of the British Ornithologists’ Union and the author of A Dictionary of Birds (1896), a paragon of erudition that was said by Coues (1897) to be “by far and away the best book ever written about birds.” Little has happened in the past 80 years to render this appraisal any less valid. Newton had a par- ticular interest in the Great Auk and, even though quite lame, visited a num- ber of its rugged former breeding places. He wrote several papers on the bird and was engaged in further work on the subject at the time of his death (Hudleston, 1909; Wollaston, 1921). It seems especially appropriate that the present species should bear his name. Diagnosis and description.—Pinguinis alfrednewtoni is separable trom 692 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON | Fig. 1. Right ulnae of Pinguinis. P. alfrednewtoni holotype (USNM 193334) on left in each pair, P. impennis on right: A, External view; B, Internal view; C, Palmar view; D, Proximal view. A-—C natural size; D 1.5~x. the Mancallinae by its proportionately longer ulna (75% of femur length vs. less than 55% in Mancalla) and by the characters of the humerus enu- merated by Howard (1966:3). Within the Alcinae it is referable to Pin- guinis by having the ulna shorter than the femur. The six fossil specimens all fall within the range of size variation of P. impennis. The ulna of P. alfrednewtoni differs from that of P. impennis in having VOLUME 90, NUMBER 3 693 the shaft more curved and decidedly less flattened, with the trailing edge more rounded. In proximal view the internal cotyla is wider and more rounded; the external cotyla is wider but not as deep as in P. impennis and does not extend as far palmarly; the olecranon is thicker, protrudes more anconally, does not curve medially at the tip, and is less sharply demarcated from the cotylae than in P. impennis by virtue of having the groove between it and the external cotyla much shallower. In external view the surface of the external cotyla is narrower and the distal tendinal groove is more nearly perpendicular (c. 70°) to the long axis of the shaft than in P. im- pennis (c. 45°). In palmar view the proximal radial depression is much narrower, the crest of the intermuscular line is less distinct, the carpal tuberosity and distal tendinal groove are wider and the external lip of the latter less developed than in P. impennis. In internal view the very nar- row and elongate attachment of the anterior articular ligament, characteris- tic of Pinguinis, is situated much more proximally, being nearly confluent with the internal lip of the internal cotyla, and having a distinct pit at its proximal extremity, whereas in P. impennis this attachment is widely separated from the internal cotyla and the pit is absent. The shaft of the humerus of P. alfrednewtoni, like that of the ulna, is less compressed than in P. impennis. This is best illustrated in the cross-section (Fig. 2d) of the better preserved of the two paratypic humeri (USNM 179226), where it is seen that towards the proximal end the shaft in P. alfrednewtoni is much more rounded, the medullary cavity is larger and the wall of peripheral bone much thicker. In P. impennis there is con- siderable variation in the degree of flattening of the shaft and particularly in the extent of the development of the crest along the proximo-anconal edge; certain individuals do not differ quite as strikingly as the one shown in Fig. 2d, while others differ more. Yet in none is the proximal end of the shaft as heavy and rounded as in P. alfrednewtoni. By contrast, the dis- tal portion of the shaft, as seen in the poorly preserved distal fragment USNM 192497, appears to be less rounded, with thinner walls, than in P. impennis. Other differences between the humeri of P. alfrednewtoni and P. impennis are more subtle. In the former, the ectepicondylar prominence is more pointed and projects farther from the shaft, the attachment of the anterior articular ligament is smaller and does not extend as far proximally, the tricipital grooves are shallower, and the internal condyle in distal view is narrower. The single available femur of P. alfrednewtoni is rather worn, but in spite of this abrasion the head appears to have been markedly smaller than in P. impennis. The rotular groove appears to be somewhat wider and shallower, but other consistent differences from P. impennis are not pres- ently detectable. 694 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 2. Left humeri of Pinguinis. P. alfrednewtoni (USNM 179226) on left in each pair (above in D), P. impennis on right (below in D): A, Internal view; B, Anconal view; C, External view; D, Cross section through proximal portion of shaft in proximal view. A-C natural size; D 1.5~x. Although quite worn, the single fragment of tibiotarsus of P. alfrednew- toni still shows some distinctions from its counterpart in P. impennis. Like that species, the tendinal bridge is not ossified, but the internal ligamental prominence is better developed and the groove on the antero-medial por- VOLUME 90, NUMBER 3 695 Fig. 3. Hindlimb elements of Pinguinis: A, Distal end of right tibiotarsus of P. alfrednewtoni (USNM 193101); B, Left femur of P. alfrednewtoni (USNM 206362) on left and P. impennis on right; C, Right tarsometatarsus of P. alfrednewtoni (USNM 179277) on left and P. impennis on right. All figures natural size. tion of the shaft just proximal to the internal condyle is deeper. The most striking difference is seen in the medial view of the internal condyle; in P. alfrednewtoni the prominent postero-distal expansion of the rim ap- pears to be lacking or greatly reduced, whereas it is well developed in P. impennis, causing the margin of the condyle to be distinctly notched. The fragment of tarsometatarsus of P. alfrednewtoni is also worn and is porous proximally so as to suggest that the specimen may have been from an immature bird. In anterior view the two portions of the tubercle for M. tibialis anticus are united and are of about equal distal extent, whereas in P. impennis the two portions are more elongate and are separated, the lateral one extending farther distally and being well separated from the lateral proximal foramen. In proximal view the internal cotyla is deeper, does not extend as far medially, and has the postero-lateral mar- gin more thickened and elevated than in P. impennis. The hypotarsus, as in P. impennis, consists of three calcaneal ridges and two grooves, but the internal ridge is thinner, the middle ridge is considerably thicker and protrudes farther posteriorly, and the external ridge is much more dis- tinct than in P. impennis. Remarks.—Despite its considerably greater age, Pinguinis alfrednewtoni 696 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON is remarkably similar in size and proportions to Recent P. impennis. The differences in the hindlimb of the fossil species are not obviously explained, but those seen in the wing elements indicate that the fossil form was slightly less specialized for wing-propelled locomotion than its Recent counterpart. This is evidenced mainly by the lesser degree of flattening of the humerus and ulna. Although a relatively minor feature, the angle of the distal tendinal groove of the ulna is not without significance. That this groove is more nearly parallel to the shaft of the ulna in P. impennis probably indicates a decreased ability to flex the distal portions of the wing, a condition that would accord with increased specialization of the wing as a rigid paddle. It seems likely that P. alfrednewtoni was the direct ancestor of P. impennis. The only evidence possibly contrary to such a hypothesis is that the con- formation of the olecranal area in P. impennis appears more similar to that of typical volant alcids than to P. alfrednewtoni. It is otherwise noteworthy that so little morphological change took place in the Pinguinis lineage during the 5 million years or so since the remains of P. alfrednewtoni were deposited. In describing the new genus and species Australca grandis from the Middle Pliocene Bone Valley Formation in Florida, Brodkorb (1955) sug- gested that the supposedly reduced wing elements of this form indicated that it was approaching flightlessness and may have been ancestral to Pinguinis. It is now known that Brodkorb’s A. grandis is actually a com- posite of at least two typically volant species of alcids (Olson and Wetmore, in press). Cranial material from Lee Creek referable to Australca appears to be morphologically intermediate between Pinguinis and the Recent genus Alca but the wing elements show no tendency towards the specializations of Pinguinis. That Pinguinis was a contemporary of Australca at Lee Creek, and occurs earlier in the fossil record than the Bone Valley specimens of A. grandis, removes any possibility that A. grandis was ancestral to Pin- guinis. The great scarcity of Pinguinis alfrednewtoni in the Lee Creek deposits, where thousands of alcid bones have been recovered, is probably due to one of two factors. Either the species did not occur regularly at the lati- tude of North Carolina, in which case its principal range was probably farther north, or it inhabited areas closer to shore than the other Lee Creek alcids. Judging from the fossil avifauna as a whole, the environment at Lee Creek during the early Pliocene was most likely open ocean at a fair distance from land (Olson and Wetmore, op. cit.) Acknowledgments For helpful comments on the manuscript I am grateful to John Farrand, Jr., Hildegarde Howard and Clayton E. Ray. I am also indebted to Thomas VOLUME 90. NUMBER 3 697 G. Gibson for assessing matrix samples and to Victor E. Krantz for the photography. Literature Cited Brodkorb, P. 1955. The avifauna of the Bone Valley Formation. Florida Geological Survey, Report of Investigations No. 14:1—57. —. 1967. Catalogue of fossil birds: Part 3 (Ralliformes, Ichthyornithiformes, Charadriiformes). Bulletin of the Florida State Museum, Biological Sciences 11:99-220. Coues, E. 1897. [Review of] Newton’s Dictionary of Birds: Part IV. Auk 14: 234-244, Greenway, J. C., Jr. 1958. Extinct and vanishing birds of the world. Special Publica- tion No. 13. American Committee for International Wild Life Protection: New Yorke 8) pp: Howard, H. 1966. A possible ancestor of the Lucas Auk (Family Mancallidae) from the Tertiary of Orange County, California. Los Angeles County Museum Con- tributions in Science 101:1-8. Hudleston, W. H. 1909. Professor Alfred Newton. Ibis, series 9, volume 2, Jubilee Supplement: 107-116. Lucas, F. A. 1890. The expedition to Funk Island, with observations upon the his- tory and anatomy of the Great Auk. Report of the United States National Museum for 1887—88:493-529. Newton, A. 1896. A dictionary of birds. Adam and Charles Black: London. 1,088 pp. Olson, S. L. 1973. Evolution of the rails of the South Atlantic Islands (Aves: Rallidae). Smithsonian Contributions to Zoology 152:1-53. , and A. Wetmore. In press. a ye oyeL YMOIS [eIIUL UBY) Ssol A[JURIIFLUBIS 44 co'0. «> «Cod:s«Ye oFeL YWMOIS [eITUL UPY} Sso| APJUROIFIUBIS , ‘Ioyeois 10 Sogo] GE = N ‘A'S = (4A/UUL) SUBOUT o1e Soe YMOIL , Se = LE Ee LEU = 69S OL6I 0G0 + OL'E OL6I-6961 960 + CE x96 0 + 68°C 4VG0 + ELE 6961-8961 ce0 = 69'S «860 + OVE VCO + 98'7F 8961-LI6I CoO + 6TE V60 = SBT L961 «x60 + 668 cc 0 + 997 O70 + 08'S LO6I-996I 8T0 + 66T 6G'0 + 889 9961 LEO = STS 2960 + GGG 9961-S96I ti SS es eee eee ee a ee eae PL yeipeng ZI yeIpeng GG JeIpEnd IT ye1peno 6 yeIpendg Q yeIpeng Ivok Dsladsuod pYyawivdoyjuvX sIsUaLOUDG DYaULIDdOpNas J (1A /UIUT) SoYCI YZMOAL) os ee ee ee ee Eee ‘Tey Surjeis0zUIsIp 10J poyepMoyeo yey} st Avie Yowe UT po}jsl] Fe YIAoIs Yse] VY, ‘eoUsdsoues Yyoeoidde Tey} se psiadsuod DyawivdoyjupKX PUe SisuUaLOUll}DG pDyawuindopnasg SUeYoI] OY} OF sozet YIMOIs oqo[-e[suls [enuue Ul sosueyy “S FGPL 712 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Figs. 19-22. Apothecia development in a growth series for Xanthoparmelia conspersa. Some apothecia grow faster than others, but the largest apothecia do not change in size. Scale in Fig. 17 for all photos in mm. Fig. 19. 4 December, 1965. Fig. 20. 6 January, 1967. Fig. 21. 16 February, 1968. Fig. 22. 15 November, 1968. to development of specific lichen structures, the kind of information that would be of value in recognizing growth phases in lichens and interpret- ing life history data in terms of population dynamics. Sernander-Du Rietz (1957) conducted a study on the factors influencing apothecia initiation and development in Parmelia tiliacea in Fennoscandia, and found that clima- tological factors, particularly high temperatures and precipitation, cor- related well with apothecia development. She found also that severe cold temperatures often killed microapothecia. Figures 19-22 illustrate the super- ficial development over a 3-year period of several selected apothecia on a mature thallus of X. conspersa from Plummers Island. Most apothecia grow slowly and consistently to a specific maximum size, but the growth of some apothecia appears to be halted shortly after initiation. This is particularly true of microapothecia located close to larger apothecia. Ad- ditional studies of this nature should be done to learn how apothecial de- velopment correlates with thallus age and size and how growth rate of the thallus is influenced by the initiation of apothecia development during the life history of a lichen thallus. VOLUME 90, NUMBER 3 713 Figs. 23-28. Successive photographs of Xanthoparmelia conspersa taken at Plummers Island. The two light circles are drill holes used for relocation of the colony. Line drawn for viewer orientation. Scale in mm. Fig. 23. 20 March, 1966. Fig. 24. 18 September, 1966. Our studies of growth phases have uncovered significant life cycle pat- terns among the lichens observed on Plummers Island. We believe that additional studies are required, particularly in view of the increasing number of studies of lichen distribution in relation to long-term, chronic air pollution (e.g., automobile exhaust). If lichen growth studies continue to show determinant life cycles for many species of lichens, many of the conclusions of lichen-air pollution studies which show gradual changes 714 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON in lichen distribution over long periods of time will have to be reinterpreted. Since so little is known about the “typical” responses of lichens to en- vironmental factors over long periods of time, we suggest that it is unwise to read too much into uncritical short-term lichen distribution studies which make use of presence-absence data or growth rate data without adequate controls. We would hope that lichenologists will feel compelled to provide answers to basic questions concerning life cycle patterns of lichens, in- cluding demographic patterns. Until these questions are answered more critically, it is hazardous to make too many assumptions regarding lichens as “indicators” of low-level air pollution. Growth Rates and Environmental Factors The influence of microclimatological factors (e.g., light, relative humidity, cloud cover, wind, etc.) on lichen growth is not well documented. Haus- man (1948) claimed that high growth rates for Parmelia centrifuga in New England correlated with high cloud cover. Karenlampi (1971) investigated the influence of several climatic factors on growth of various Cladonia ssp. during a summer in Finland. He found that lichen growth was influenced mostly by rainfall and to a lesser degree by temperature and light. Nienburg (1919) reported that heavy rains in Germany in 1899 may have been responsible for high growth rates there. Also, Andreev (1954) and Barashkova (1961) have stated that areas of high humidity will support high growth, and Showman (1976) found that Parmelia caperata grew better on more consistently moist sides of a tree than on exposures which dried out quickly. Miller’s (1966) study of lichen growth and thallus wet- ness and Karenlampi’s (1971) study of fruticose lichen growth rates are among the few which make use of microclimatological data collected at the level of the plants themselves. Weather summaries collected many miles from the study site are not sufficiently exact to be used in lichen growth studies without more precise corroborating evidence. There is also a need for statistical analysis of growth data which allows broad inferences to be drawn from large amounts of data. Consequently, the use of com- puters to calculate growth rates and to determine statistically significant relationships between growth and various environmental variables is be- coming more frequent (Karenlampi, 1971). A summary of Plummers Island weather variables for the years 1966— 1969 (Fig. 29) shows maximum-minimum temperatures and total precipi- tation data collected at the growth study sites. We computed average precipitation (total precipitation in period/days of precipitation in period) and percent cloudy days (number days in period with 60% of greater cloud cover/total number days in period x 100) for each period from National Airport weather summaries. During the four years of weather sampling, temperature patterns were relatively predictable with maximum VOLUME 90, NUMBER 3 715 Fig. 25. 13 August, 1967. Fig. 26. 15 November, 1968. (See legend for Figs. 23— 28.) temperatures approching 40°C in the summer and minimum temperatures often -15°C in the winter. Precipitation, however, was not constant. Total precipitation in 1968, for example, was much lower than in 1969. Analysis of cloud cover data revealed no seasonal pattern of cloud cover from 1966-1969. Scatter plots of daily lichen single-lobe growth rates vs. various weather variables were computer-generated for Pseudoparmelia baltimorensis and Xanthoparmelia conspersa. Because growth rates of X. conspersa_ ex- hibited little seasonal or annual variation, no weather-related growth 716 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 27. 9 April, 1969. Fig. 28. 17 November, 1969. (See legend for Figs. 23-28.) patterns were discernible for this species. However, relationships were apparent between growth rates of P. baltimorensis and_ cer- tain weather variables. Scatter diagrams of radial growth vs. high tem- perature (Fig. 30) and low temperature (Fig. 31) in each sampling period show that optimum growth for P. baltimorensis occurs during periods in which the maximum temperature is 25-30°C and the minimum temperature is around 0°C. A glance at Fig. 29 shows that these temperature condi- tions occur on Plummers Island in the early spring. It is possible that during this time, increasing irradiation coupled with minimum canopy cover pro- duce relatively high temperatures during the day and low temperatures at VOLUME 90, NUMBER 3 ih aS) a. | B50! -10 ; | Lil da hl heat elit el 1966 1967 1968 1969 Fig. 29. Precipitation (vertical lines) and maximum-minimum temperatures (hori- zontal lines) recorded at Plummers Island for each sampling interval from 1966 to 1969. night because of radiation loss. As both maximum and minimum tem- peratures increase during the summer, lichen growth rate decreases. It appears from the curves of lichen growth vs. temperature (Figs. 30 and 31) that temperatures rarely exceed critical limits for growth of P. balti- morensis. Growth of this lichen species, therefore, is possible at varying rates throughout the year on Plummers Island. Scatter diagrams of P. baltimorensis growth vs. total precipitation (Fig. 32) and average precipitation (Fig. 33) in each sampling period indicate that lichen growth can actually be inhibited in periods of high precipitation. High growth of P. baltimorensis may conversely occur under conditions of low precipitation; optimum growth takes place when average precipita- tion (total precipitation/days of precipitation in period) is between 0.5-1.0 cm/day. Karenlampi (1971) detected high levels of linear correlation be- tween growth of fruticose lichens and rainfall (mm/day) in Finland, al- though rainfall data used in his study appeared to range from only 0-4 mm/day, considerably less than the maximum rainfall observed on Plum- mers Island. Within this narrow range of rainfall values, our growth data also show a high correlation. However, there is a considerable decline in growth once optimum rainfall is exceeded. The graph of growth rate for P. baltimorensis vs. percent cloudy days (Fig. 34) shows that optimum growth occurs during sampling periods which are cloudy (60% cloud cover or more) more than half of the time. Growth rate apparently decreases after values of percent cloudy days reach 75% 718 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON PSEUDOPARMELIA BALTIMORENSIS 0.14 x 0.13 0.12 0.11 0.10 x x x 0.09 x a x 2 0.08 = x >= -07 = x a x Po : x x= 0.06 x x hex = ro) x Xx a x x x i eae é x x 0.04 e : x xi xX x X< 9 x x ae x x x ; x 0.03 ee SOK X ~ x x ix x * x 3. 10 X x x x x * x K X x 0.02 * Ox ex x x xox & x ox oe x xx x x x % 4 x x IWS x x x x be Bok 9K xx x 0.01 X »& x Kes ‘ ¥ x =X x FOS % O06 * Y x x x RY ew xX Baw KK OR x , ay ee ° x X x 0.00 bh -o- =. ee eos 8 Q. . 10. 15. 0 30. - 40. 4s. QO. HIGH TEMP (C) Fig. 30. Daily radial single-lobe growth rate of Pseudoparmelia baltimorensis vs. maximum temperature in each sampling period at Plummers Island from 1966 to 1969. or more, although growth is active even when every day during the sampling period exhibits 60% or greater cloud cover. There is greater inhibition of P. baltimorensis growth during periods of little cloudiness than during periods when almost every day in the sampling period exhibits high cloud cover. This would indicate that possibly high irradiation is a more limiting environmental factor for P. baltimorensis than cloudiness. Hausman (1948) believed that high growth rates of Parmelia centrifuga in New England were due to high cloud cover. Our data would tend to support this view, with the understanding that other weather variables (temperature, relative hu- midity, irradiation) also change in direct proportion to the degree of cloudiness. Stepwise multiple regressions of mean daily growth per sampling period VOLUME 90, NUMBER 3 AS PSEUDOPARMELIA BALTIMORENSIS 0.14 x x 0.13 0.12 Aah 0.10 x x x 0.09 x « x 2 9.08 x x x x= w0.07 x x = x x x x @ . x = 0.06 x x MY x 3 ie x x x © 0.05 xX KY x x x x x MY x x 0.04 x x XK Zs 5% 0) i x x x x x MX 0.03 ain . x x x Xx x x K HL KX x x Xx § x xX x x x 0.02 x nee aX Ge ‘ sec Bx x Nakx oe J 0.01 x x X RX aXe : % xx Rx x x x x % 20K s x x ¥ xe x xX SS x I A ue ex 0.00 -30. -20. -10. ° 20. LOW TEMP (C) Fig. 31. Daily radial single-lobe growth rate of Pseudoparmelia baltimorensis vs. minimum temperature in each sampling period at Plummers Island from 1966 to 1969. on numerous weather variables were computed to determine which weather variables or combinations of weather variables explained the greatest amount of the variance in growth exhibited by Pseudoparmelia baltimorensis. Be- cause we did not collect data on irradiation, aspect, or non-weather factors, we can make no determination about the relative importance of these factors even though other studies have shown they probably are im- portant (Haukulinen, 1966; Jones and Platt, 1969; Karenlampi, 1971). A regression equation using the variables high temperature, total precipitation and percent cloudy days was found to explain 22% of the variation in lichen growth rate for P. baltimorensis (r = 0.466, B significantly different from 0 at p < 0.05). Of these three weather variables, percent cloudy days explained the greatest amount of variation (14%, r = 0.376). The gen- 720 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON PSEUDGPARMELIA BALTIMORENSIS 0.09 x GROWTH RATE (MM/DAY } x x x x x x x x x 0. 2s. 30. TOTAL PRECIP (CM) Fig. 32. Daily radial single-lobe growth rate of Pseudoparmelia baltimorensis vs. total precipitation in each sampling period at Plummers Island from 1966 to 1969. erally low correlation between weather variables and lichen growth was thought to be due to the non-linear nature of the relationship. We re- gressed logio, In, x“, and x? transformations of the growth data on the various weather variables in order to determine if these transformations rectified the growth data. We found that the regression of (mean lichen growth rate)? on percent cloud cover explained 19% of the growth variance (r = 0.436, significantly different from 0 at p < 0.05). This would indicate that possibly radial growth measurements contain less information than area measurements, but the increase in predictability was not overwhelm- ing. Addition of the other two variables (total precipitation and high tem- perature) to the regression equation did not significantly increase the pre- dictability. We concluded that percent cloudy days is most useful as a a VOLUME 90, NUMBER 3 721 4f SEUDOPARMELTA BALTIMORENSIS GROWTH RATE (MM/ODAY) 2.0 2.5 3.0 . . . -0 AVERAGE PRECIP (CM) Fig. 33. Daily radial single-lobe growth rate of Pseudoparmelia baltimorensis vs. average precipitation (total precipitation in each sampling period/days of precipitation in each sampling period) at Plummers Island from 1966 to 1969. predictor of lichen growth. However, none of the weather variables we measured was especially useful in this regard, since a total of only 22% of the growth rate variance is explained by any weather variable or combina- tion of variables. Karenlampi (1971) found high correlation between weather variables and fruticose lichen growth over short time periods. We believe this may be a result of measuring lichen growth only during favorable periods, inasmuch as lichen growth can be expected to respond more strongly to changes in weather when weather conditions are most favorable for growth. Additionally, Karenlampi’s (1971) study made use of irradiation data, which the present study does not include. This factor may contribute to lichen growth on Plummers Island as it does in Fin- 722 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON PSEUDOPARMELIA BALTIMGRENSIS 0.09 GROWTH RATE (MM/DAY ) x x x x x 0.04 x . x Byes ee x x as * 0.03 x % x x x x ZS ‘ x x x % XE 0.02 x %%x x x x x x % x & x ig" x x x XE 0-01 ™ x x Se x * x x x xe x Kx x x ae We % x% x x . 0. . 60. CLOUDY DAYS (PERCENT) Fig. 34. Daily radial single-lobe growth rate of Pseudoparmelia baltimorensis vs. percent cloudy days (number days of 60% cloud cover or more/total number days in sampling period x 100) at Plummers Island from 1966 to 1969. land, but we would expect that it would not increase correlation to the levels observed by Karenlampi. Additional long-term studies of the influence of weather on other lichen species in other geographic areas will allow comparisons to be made with the few studies presently available. At the present time, lichen growth appears to correlate with certain weather variables. However, we are still unable to predict changes in lichen growth over long periods of time with certainty. Conclusions A twelve-year growth study of saxicolous lichens from Plummers Island, Maryland, revealed numerous differences in annual and daily radial growth VOLUME 90, NUMBER 3 723 rates between various lichen species and variously-aged lichens of the same species. Growth rates were tested for significant correlations with weather variables. The major findings can be listed as follows: (1). The various large-lobed Parmelias exhibited the highest growth rates (around 5 mm/yr), followed by Xanthoparmelia conspersa, Physcia millegrana, and various crustose lichen species. This pattern illustrates a possible relationship between thallus lobe morphology and growth rate which requires further study. (2). There were significant differences in growth rates between juvenile and mature thalli of Pseudoparmelia baltimorensis and X. conspersa, the juvenile thalli growing at reduced rates. There were also seasonal vari- ations in growth from year-to-year for each species studied, although dif- ferences between species occurred consistently regardless of annual or seasonal variations in growth. (3). There was a great deal of variation in growth between lobes of the same thallus, indicating that large sample sizes are necessary in order to generalize from growth rate data. (4). Differences in growth rate patterns appeared to be related in part to senescence, with measurable differences in growth between thalli before and after disintegration. Growth rates of disintegrating P. baltimorensis thalli were significantly reduced from growth rates before disintegration. Xanthoparmelia conspersa thalli exhibited no significant differences in growth before or after thallus disintegration, possibly because reduced growth rates made detection of patterns more difficult for this species. Thallus disintegration appeared to be a rejuvenation process rather than a true senescence, inasmuch as growth rates after disintegration approximated those of juvenile thalli. No lichen thalli of the dominant species photo- graphed on Plummers Island were observed at diameters much larger than 15 cm. (5). Apothecia development was observed for X. conspersa. No correla- tions between apothecia development and other factors were possible due to a paucity of data. However, there appeared to be a maximum diam- eter for apothecia of X. conspersa, and microapothecia were observed to increase in size slowly or not at all, as if inhibited by adjacent larger apothecia. (6). Scatter diagrams of lichen daily growth rate vs. various weather variables showed that optimum growth of Pseudoparmelia baltimorensis oc- curred on Plumers Island during sampling periods in which the maximum temperature was around 25-30°C, the minimum temperature around 0°C, the precipitation around 0.5-1.0 cm/sampling period, and the cloud cover 60% or more for at least half the sampling period. All the lichen growth vs. weather scatter plots showed lower lichen values above and below these optimum weather conditions; however, there were no weather conditions 724 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON measured on Plummers Island which consistently correlated with zero lichen growth. Xanthoparmelia conspersa exhibited no detectable rela- tionships between growth rate and weather variables, a result probably due to the fact that growth rate values for this species exhibited little change from one sampling period to another. (7). Stepwise multiple regressions of lichen growth of P. baltimorensis on weather variables (high temperature, total precipitation, cloud cover) re- vealed that a total of 22% of the lichen growth variance could be ex- plained by any combination of weather factors. The cloud cover variable consistently explained the greatest amount of the lichen growth variance. Squaring the lichen growth dependent variable resulted in a somewhat higher correlation with weather variables, which may be due to the fact that radial growth data were used rather than area data. Literature Cited Andreev, V. N. 1954. Growth of forage lichens and methods of improving it. Geo- botanika 9:11-74. [In Russian.] Armstrong, R. A. 1973. Seasonal growth and growth rate-colony size relationships in six species of saxicolous lichens. New Phytol. 72:1023—-1030. —. 1974. Growth phases in the life of a lichen thallus. New Phytol. 73:913-918. —. 1976. Studies on the growth rates of lichens. Pp. 309-322 in D. H. Brown, D. L. Hawksworth and R. H. Bailey (eds.), “Lichenology: Progress and Prob- lems.” Academic Press, London. Barashkova, E. A. 1961. Cladonia rangiferina (L.) Web. in the conditions of the district of Murmansk. Bot. Zur. 46:410-414. [In Finnish.] Beschel, R. 1958. Flechtenvereine der Stadte, Stadflechten und ihr Wachstum. Ber. Naturwiss. -Med. Ver. Innsbruck 52:1-—158. Broadhead, E., and I. W. B. Thornton. 1955. Elipsocus melachlani feeding on lichens. Oikos 6:1-50. Brodo, I. M. 1965. Studies of growth rates of corticolous lichens on Long Island, New York. Bryologist 68:451—456. Coker, P. D. 1967. Damage to lichens by gastropods. Lichenologist 3:428-429. Hakulinen, R. 1966. Uber die Wachstumgeschwindigheit einiger Laubflechten. Ann. Bot. Fenn. 3:167-179. Hale, M. E., Jr. 1967. The Biology of Lichens. Edward Amold, London. 176 p. 1970. Single-lobe growth rate patterns in Parmelia caperata. Bryologist 73: 72-81. ——. 1972. Natural History of Plummers Island, Maryland. XXI. Infestation of the lichen Parmelia baltimorensis Gyel. & For. by Hypogastrura packardi Folsom (Collembola). Proc. Biol. Soc. Wash. 85:287-296. ——.. 1973. Growth. Pp. 473-492 in V. Ahmadjian and M. E. Hale, Jr. (eds.), “The Lichens.” Academic Press, N.Y. Hausman, E. H. 1948. Measurements of the annual growth-rate of two species of rock lichens. Bull. Torr. Bot. Club 75:116-117. Jones, J. M., and R. B. Platt. 1969. Effects of ionizing radiation, climate and nutrition on growth and structure of a lichen Parmelia conspersa (Ach.) Ach. Radioecol. Concent. Processes, Proc. Int. Symp., 2nd, 1969, pp. 111-119. VOLUME 90, NUMBER 3 725 Karenlampi, L. 1970. Morphological analysis of growth and productivity of the lichen Cladonia alpestris. Rep. Kevo Subarctic Res. Sta. 7:9-15. —. 1971. Studies on the relative growth rate of some fruticose lichens. Rep. Kevo Subarctic Res. Sta. 7:33-39. Laundon, J. R. 1971. Lichen communities destroyed by psocids. Lichenologist 5: ee Miller, A. G. 1966. Lichen growth and species composition in relation to duration of thallus wetness. B.S. Thesis, Queen’s University, Lingston. Nienburg, W. 1919. Studien zur Biologie der Flecten. I. II. III. Z. Bot. 11:1-38. Peake, J. F., and P. W. James. 1967. Lichens and mollusca. Lichenologist 3:425-— 428. Phillips, H. C. 1963. Growth rate of Parmelia isidiosa (Mull. Arg.) Hale. Jour. Tenn. Acad. Sci. 38:95-96. Rydzak, J. 1961. Investigations on the growth rate of lichens. Ann. Univ. Mariae Curie-Sklodowska, Sect. C. 16:1-15. Sernander-Du Rietz, Greta. 1957. Om yttre faktorers inverkan pa apothecie-bildningen hos Parmelia tiliacea. Svensk Botanisk Tidskrift 51:454—-488. Showman, R. E. 1976. Seasonal growth of Parmelia caperata. Bryologist 79:360- 363. Sowter, F. A. 1971. Mites (Acari) and lichens. Lichenologist 5:176. Footnote ‘The preceding number in this series is published in Proceedings of the Biological Society of Washington, Vol. 90, No. 3, pp. 615-647. Publication costs of the present number have been defrayed by the Washington Biologists’ Field Club to promote its primary objective of research on the fauna and flora of Plummers Island and adjacent areas. (JTL) Ohio State University, Columbus, Ohio 43210; and (MEH, Jr.) Smithsonian Institution, Washington, D.C. 20560. PROC. BIOL. SOC. WASH. 90(3), pp. 726-734 A GILL-INHABITING NEW GENUS AND SPECIES OF THE BRANCHIOBDELLIDA (ANNELIDA: CLITELLATA) Perry C. Holt Abstract—Holt, Perry C., Department of Biology and Center for Syste- matics Collections, Virginia Polytechnic Institute and State University, Blacksburg, Virginia 24061.—Adenodrilus hemophagus, from Douglas County, Oregon, U.S.A., is described, its affinities discussed, and evidence of its way of life presented. The first formally described branchiobdellid, Branchiobdella astaci Odier, 1823, apparently is an inhabitant of the gill chambers of the host (Odier, 1823:75). Though there is little doubt that some European branchiobdellids are adapted to life in branchial chambers, the taxonomy of European branchiobdellids appears to be in a state of confusion. Pop (1965) was undoubtedly correct in reducing many specific names to synonymy, but I doubt that there are only four species of branchiobdellids in Europe and I have no idea how many are true ectoparasites that feed on the host's branchiae. In Asia, Yamaguchi (1934:216) reported that two species of “Stephano- drilus’ (= Cirrodrilus) are found in the gill chambers of the host and two others of this genus are found in both the gill chambers and on the body surface. The knowledge of the gill-inhabiting branchiobdellids of North America is almost as scanty. Bdellodrilus illuminatus (Moore, 1893) is common and known to live in the branchial chambers and feed on the blood of the host sucked from filaments of the gills. Cambarincola branchiophilus Holt, 1954, also inhabits the gill chambers of crayfishes and some species, e.g., Cam- barincola demissus Hoffman, 1963, are suspected of doing so. No effort to study gill-inhabiting branchiobdellids as such has been undertaken. In part, this is because the method of collecting, satisfactory for those worms that live on the exposed surfaces of the host, is not so good for the branchial forms; they tend to be excessively shrunken and contorted by the alcohol- formalin fixative I use. In addition the usual method of preparing the ani- mals for study (dehydrating in alcohol, clearing in clove oil and mounting in balsam) exacerbates this difficulty. The specimens described herein, nonetheless, were prepared by these methods and, with the aid of dissections, a satisfactory account of their structure was made possible. In other respects, my usual methods were used (Holt, 1960), and all drawings are oriented with the anterior of the animal or part thereof to the reader's right. VOLUME 90, NUMBER 3 727 It seems appropriate, therefore, to call attention to this population of branchiobdellids, representing a new genus, from Douglas County, Oregon, since previous records of gill-inhabiting North American branchiobdellids are so few. I wish to acknowledge the aid of my wife and daughter in collecting; Dr. Horton H. Hobbs, Jr., for identifying the hosts; Dr. Marian Pettibone for her many kindnesses in cataloging type-specimens and reviewing manu- scripts. Adenodrilus, new genus Type-species.—Adenodrilus hemophagus, here designated. Diagnosis.—Relatively large worms, unpaired anterior nephridiopore; body terete; body-wall muscles thin; head markedly less in diameter than body segments; sucker subequal to head in diameter; peristomium entire to faintly lobed; jaws triangular with prominent lateral flanges; spermiducal gland with vasa deferentia entering ectad to ental end, often expanded; no prostrate or prostatic protuberance; ejaculatory duct long, thick, heavily muscular; bursa elongate pyriform; penis muscular; spermatheca with spermathecal bursa, long ectal duct, expanded bulb, long glandular ental process. Etymology.—Adenos, G., gland and drilos, G.—“gland-worm.” Affinities—The most recent attempt to develop a scheme of relation- ships among branchiobdellid genera (Holt, 1969) was proposed rather timidly and the discovery of Adenodrilus increases the difficulties of de- vising satisfactory theories of phylogenetic relationships within the order. For now, I can only postulate that the adaptive radiation of the group occurred early in its history and that considerable parallelism or con- vergence or both characterized its course. The best that can be done is to briefly consider a few genera that share important features with Adeno- drilus. Four genera were placed in the Cirrodrilus-lineage (Holt, 1969:197) and Adenodrilus fits nicely with these (Cirrodrilus Pierontoni, 1905; Branchiob- della Odier, 1823; Xironogiton Ellis, 1919; Ankyrodrilus Holt, 1965) ex- cept that all have two anterior nephridiopores. Cronodrilus Holt, 1968b, possesses only one anterior nephridiopore and the vasa deferentia enter the spermiducal gland ectad to its ental end as in Adenodrilus. But the penis of the latter is muscular, instead of the eversible, non-cellular, cuticu- lar, tubular one of Cronodrilus and is doubtfully eversible (see below). Among the genera characterized by a spermiducal gland in which the vasa deferen- tia enter the gland ectad to its ental end, Cirrodrilus appears to be most 728 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON nearly similar to Adenodrilus. Indeed, except for the two anterior nephrid- iopores of Cirrodrilus inukaii (Yamaguchi, 1934) and the different structure of the jaws, C. inukaii and the species of Adenodrilus described below could be considered as congeneric. Yamaguchi (1934 and previous papers) did not designate a type-species for his genus “Stephanodrilus” (= Cirrodrilus Pierantoni 1905). But both C. inukaii (the only species adequately illustrated) and C. cirratus have un- paired anterior nephridiopores. Adenodrilus, thus, shares only two im- portant features, the manner of entry of the vasa deferentia into the spermi- ducal gland and the basic structure of the bursa and penis with Cirrodrilus. So, the unexpected combination of an unpaired anterior nephridiopore and an ectal entry of the vasa deferentia into the spermiducal gland, leaves me with only one dubious choice in an attempt to place Adenodrilus in my former scheme (Holt, 1969: Figure 2). It seems to belong to the group com- posed of Cronodrilus and Bdellodrilus Moore, 1895, except for its muscular and possibly non-eversible penis and, on the assumption that muscular, pro- trusible penes have evolved from eversible, cuticular ones several times in the radiation of the branchiobdellids, I must, for now, leave Adenodrilus there. Geographical and other remarks.—No great significance should be at- tached to the geographical location of Adenodrilus. I have previously commented upon the relationships of the American and Asian branchiob- dellid faunas (Holt, 1968a; 1969) and postulated an ancient origin for the genera of the order (Holt, 1968a; 1969; 1977). The discovery of Adeno- drilus is only one of several additional bits of evidence supporting these speculations. So little is known of the ecology, physiology, feeding habits, etc., of the branchiobdellids, that the basis of my assertion that A. hemophagus feeds on blood drawn from the host’s gills, in the absence of direct observation, requires comment. There is a recognizable parasitic facies common to the truly gill-inhabiting, blood-sucking species. In Bdellodrilus illuminatus and Adenodrilus hemophagus the body-wall is thin and the glandular elements of the body-wall and the sucker, by virtue of the reduction of the muscula- ture, accentuated. The coelomic spaces seem proportionately greater in volume. The gut is filled with a liquid. These characteristics of gill-in- habiting branchiobdellids account for the difficulties of preparing speci- mens for study by the usual methods. And it is on the basis of this “para- sitic facies” and the invariable presence within the gut of specimens of A. hemophagus of a clear fluid coagulum without detritus of any sort or the remains of any other animals whatsoever that I am confident of the habitat on the host and the food of A. hemophagus: the branchial chambers and the blood of the hosts. VOLUME 90, NUMBER 3 729 Fig. 1. Adenodrilus hemophagus: Lateral view of holotype. Adenodrilus hemophagus, new species Figs. 1-5 T ype-specimens.—Holotype, USNM (Smithsonian Institution) 54640, 16 paratypes, PCH 1333 (VPI & SU Center for Systematics Collections), from Pacifastacus leniusculus klamathensis (Stimpson, 1857) taken from Elk Creek, Douglas Co., Oregon, at junction of State Road 231 and U.S. High- way 99, 20.3 km SW of Cottage Grove, by Perry C. and Virgie F. Holt, 11 July 1960. Diagnosis.—As for the genus. Etymology.—Hemos, G., blood, and phagos, G., eater. Description.—Specimens of A. hemophagus are relatively large branchiob- dellid worms. The holotype has the following approximate dimensions (in mm): total length, 5.0; greatest diameter, 1.2; head length, 0.7; head diameter, 0.5; diameter, segment I, 0.5; diameter sucker, 0.6. Five paratypes, selected at random, have the following approximate average dimensions (ranges given in parentheses): total length, 5.2 (4.0-6.6); greatest diam- eter, 1.1 (1.0-1.1); head length, 0.7 (0.6-0.8); head diameter, 0.4 (0.4-0.5); diameter, segment I, 0.5 (0.5-0.6); diameter, sucker, 0.5 (0.5-0.5). These animals are remarkably uniform in size. Most are bent in preservation into a semicircle and the variation in total length is probably an artifact of the difficulty of obtaining precise measurements with an ocular micrometer. The worms have a corpulent appearance; anterior and posterior portions are noticeably less in diameter than the mid-part of the body (Fig. 1). The body wall is remarkably thin, the longitudinal musculature is sparse and 730 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON a {' V b > 2 Cc Fig. 2. Adenodrilus hemophagus: a, Optical section through anterior nephridiopore; b, Lateral view of jaws; c, Peristomium, from specimen without peristomial lobes. From paratypes. its individual cells slender. The glandular components of the epidermis are prominent by virtue of the paucity of muscles, including those of the clitellar region (segments VI and VII). The sucker, contrary to the usual branchiobdellid condition, lacks a specialized musculature; instead it is provided with a cluster of about a dozen large unicellular glands whose ducts discharge onto its concave surface. : The head is marked externally by one annular sulcus in addition to the peristomial one. The lips of the peristomium are usually marked by in- distinct indentations into 4 upper and 2 lower lobes, but the peristomium may appear entire (Fig. 2c). Internally, there is one pharyngeal sulcus. There are no oral papillae. The anterior nephridiopore (Fig. 2a) is unpaired and the nephridial outlet duct is surrounded by densely granular cells. The body is often completely flexed into a circle in preservation, making observation of internal structures very difficult, a diffculty increased by the density of the spermatozoa which fill souiems V and VI. The overall facies of the animals is a parasitic one. The gut is expanded, as usual, in segments III-IX, its expansion in seg- ment VIII extends into segment IX. The gut is filled throughout its length with a homogeneous, non-particulate, fluid. The ganglia of the nervous system are proportionately small; the major blood vessels proportionately large. The jaws are prominent, medium dark brown, en ede in lateral aspect, but in actuality essentially U-shaped in frontal view, with large lateral flanges. Each jaw is equipped with a prominent median tooth; no lateral VOLUME 90, NUMBER 3 | 731 Fig. 3. Adenodrilus hemophagus: a, Latero-dorsal view of male reproductive sys- tem; b, Same of spermatheca. From a paratype. Abbreviations: b, bursa; ejd, ejaculatory duct; ep, ental processes of spermatheca; es, ental end of spermiducal gland; sb, spermathecal bulb; sd, spermathecal ectal duct; spb, spermathecal bursa; spg, spermi- ducal gland; vd, posterior vas deferens. teeth were observed. The dental formula is, then, the somewhat rare one Oily Ae (Ee: 2b). The vasa deferentia are thick and enter the spermiducal gland at about % the length of the latter ectad to its ental end. The spermiducal gland is often constricted (Fig. 3a; 4a) at the point of entry of the vasa deferentia and tapers ectally to its junction with the ejaculatory duct. It appears in 2 forms which are presumably dependent upon the reproductive state of the animal. Often it is rather slender, with a thick glandulo-muscular wall and with spermatozoa present in its lumen (Fig. 5). In other cases, it is greatly extended, filled with a clear fluid, and spermatozoa are absent or difficult to detect (Fig. 3a). The ejaculatory duct is unusual. Subequal to the bursa in length, it ap- pears to be composed of 3 layers of muscles: inner and outer longitudinal ones and an inner circular one (Fig. 5). Its structure suggests that it is capable of a strong pumping action. 732 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 4. Adenodrilus hemophagus: a, Dorso-lateral view of male reproductive sys- tem; b, Ventro-lateral view of spermatheca. Paratypes. The bursa is elongated pyriform in shape, broadest at the juncture of its penial sheath region and the bursal atrium. The atrium constitutes about %s of the organ and its walls taper to a narrow outlet duct. The penis is composed of dense muscle and hence, by analogy with species with similar penes, is protrusible. But in the absence of direct observation, this con- clusion is uncertain: the lumen of the penis, though obscure, is a meander- ing canal enclosed in the muscles of the organ and is undoubtedly longer than the retracted penis, suggesting that the penis is eversible (Fig. 5). The total size of the male reproductive organs is, proportionate to the diam- eter of the segment in which they lie, rather small; they extend dorsad about % of the distance to the dorsum of the body. The spermatheca is clearly differentiated into 4 distinct regions (Fig. 3b). Ectally, there is a spermathecal bursa with heavily muscular walls and a lumen expanded to form a cavity remarkably similar to the atrium of the male bursa. This is followed by a spermathecal duct with a wall composed of rather thin bands of muscle cells and a wide lumen. The wall of the spermathecal bulb is relatively thin but ensheathed in muscle, the lumen expanded as expected, but, surprisingly often with little evidence of stored spermatozoa within it. The spermatheca ends entally in a long prominent ental process, though the organ often is bent in such wise that VOLUME 90, NUMBER 3 SN LZ ) LE } = {}}) | All 733 Fig. 5. Adenodrilus hemophagus: Detail (schematic) of part of male reproductive system. Abbreviations: ba, bursal atrium; ejd, ejaculatory duct; p, penis; ps, penial sheath of bursa; spg, spermiducal gland. Paratype. the ental process is obscured (Fig. 4b). The inner wall of the ental process is composed of glandular cells, each with a long process extending to al- most the center of the lumen (Fig. 3b). Variations —The absence of lobes of the lips in some specimens, the re- markable differences in the degree of distension of the spermiducal gland and the various positions in which the reproductive organs may lie have been noted and to some extent illustrated: a somewhat unusual range of variations, but the overall correlation of parts among the specimens studied, leaves no doubt as to the conspecificity of these worms. Affinities—As for the genus. Host.—Pacifastacus leniusculus klamathensis (Stimpson, 1857). Distribution —Known only from the type-locality. Material examined.—The type-series. 734 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Literature Cited Ellis, Max M. 1919. The branchiobdellid worms in the collections of the United States National Museum, with descriptions of new genera and species. Proc. U.S. Nat. Mus. 55:241-265, pls. 10-13. Hoffman, Richard L. 1963. A revision of the North American annelid worms of the genus Cambarincola (Oligochaeta: Branchiobdellidae). Proc. U.S. Nat. Mus. 114:271-371. Holt, Perry C. 1954. A new branchiobdellid of the genus Cambarincola (Oligochaeta, Branchiobdellidae) from Virginia. Virginia Jour. Sci., N.S. 5:168—172. ——. 1960. The genus Ceratodrilus Hall (Branchiobdellidae, Oligochaeta), with the description of a new species. Virginia Jour. Sci., N.S. 11:53-77. ——. 1965. On Ankyrodrilus, a new genus of branchiobdellid worms. Virginia Jour. Sci., N.S. 16:9-21. ——. 1968a. The Branchiobdellida: epizodtic annelids. The Biologist 50:79-94. ——. 1968b. New genera and species of branchiobdellid worms (Annelida: Clitellata). Proc. Biol. Soc. Wash. 81:291-318. ——. 1969. The relationships of the branchiobdellid fauna of the southern Ap- palachians. In Holt, Perry C. (ed.). The distributional history of the southern Appalachians. Part 1: Invertebrates. Res. Div. Monogr. 1, Virginia Polytechnic Institute: 191-219. . 1977. An emendation of the genus Sathodrilus Holt 1968 (Annelida: Branchiob- dellida), with the description of four new species from the Pacific drainage of North America. Proc. Biol. Soc. Wash. 90(1):116—-131. Moore, J. Percy. 1894. On some leech-like parasites of American crayfishes. Proc. Acad. Nat. Sci. Philadelphia (for 1893):419—428. —. 1895. The anatomy of Bdellodrilus illuminatus, an American discodrilid. Jour. Morph. 10:497-541. Odier, Auguste. 1823. Mémoire sur le branchiobdelle, nouveau genre d’annelides de la famille des hirudinées. Mem. Soc. d’Hist. Nat. 1:71-78. Pop, Victor. 1965. Systematische Revision der europdischen Branchiobdelliden (Oligo- chaeta). Zool. Jb. Syst. 92:219-238. Pierantoni, Umberto. 1905. Cirrodrilus cirratus n. g. n. sp. parassita dell’ Astacus japonicus. Ann. Mus. R. Univ. Napoli 1:1-3. Yamaguchi, Hideji. 1934. Studies on Japanese Branchiobdellidae with some remarks on the classification. Jour. Fac. Sci., Hokkaido Imp. Univ. ser. VI. Zool. 3: 177-219. PROC. BIOL. SOC. WASH. 90(3), pp. 735-752 LARVAL AND EARLY POST-LARVAL STAGES OF THE WEST INDIAN SPIDER CRAB, MITHRAX SPINOSISSIMUS (LAMARCK) (DECAPODA: MAJIDAE) Anthony J. Provenzano, Jr., and W. N. Brownell Abstract—Provenzano, A. J., and W. N. Brownell; (AJP) Institute of Oceanography, Old Dominion University, Norfolk, Virginia 23508; (WNB) Department of Ichthyology, American Museum of Natural History, New York, N.Y. 10024.—Larvae of Mithrax spinosissimus, a large West Indian spider crab were reared in mass culture, but without food, from hatching to first crab stage in 5-6 days. The prezoea, two zoeal stages, the megalopa and first crab stage are described. Despite limited descriptions of Mith- racinae larvae in literature, they are reviewed and compared with those of M. spinosissimus. This is the first species of the genus for which the megalopa is known. For the mithracine crabs, no generic or subfamilial larval or megalopal characters are yet distinguishable with certainty. The family Majidae, containing approximately 900 species, is widely dis- tributed in marine waters. The vast majority of species are small and of no direct economic importance, but some species support commercial fisheries. Perhaps the largest majid crab in the tropical western Atlantic is the red spider crab, Mithrax spinosissimus (Lamarck, 1818), the known distribu- tion of which extends from the Carolinas through the Florida Keys and West Indies to Guadeloupe from shallow depths to 98 fathoms (Williams, 1965) and which we have now found in the Venezuelan archipelago of Los Roques. This crab is taken by fishermen and eaten occasionally but is not fished in large numbers because of its apparently low density. Despite its scarcity in the regular catch, the species may meet many of the criteria for desirable mariculture candidates. Unfortunately there is an almost total lack of published information concerning the biology and ecology of this crab, though Hazlett and Rittschof (1975) have studied local movements of adults. In 1976 this species was cultured in the laboratory from egg to advanced juvenile stages for purposes of evaluating its potential for Caribbean mariculture (Brownell, Provenzano, and Martinez, in press). Because of other activities at the time and the lack of completely adequate facilities, those experiments were not conducted under ideal conditions nor with in- tent to obtain adequate material for systematic studies, but a few speci- mens of each of the earliest stages were preserved. The purpose of the present paper is to describe, so far as the limited material will permit, the development of this crab from hatching through 6 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON (ey) the first post-larval stages and to compare the developmental stages with those of related majids. Methods and Materials In January 1976, a pair of large spider crabs, subsequently determined to be Mithrax spinosissimus, previously unknown from Venezuela, was brought by fishermen to the Los Roques field station, in the Los Roques Archipelago, 120 km north of La Guaira, Venezuela. The berried female was placed in a 1,000 l-capacity tank where the eggs subsequently hatched. Several thou- sand larvae were removed and placed into 3 aquaria, one of them con- taining phytoplankton but the others containing no apparent foods. All died before metamorphosis. An additional 24 larvae from the original hatching tank were placed in compartmented plastic trays containing filtered water. A few were given phytoplankton, the others were left unfed. Temperature ranged from 24-28°C, salinity 34-36%. Water was changed daily. Samples of the stages from prezoea through megalopa were preserved in 7% formalin. Specimens of early crab stages were recovered from the original hatching tank where larval development had proceeded unobserved. In subsequent months additional adult specimens were obtained and broods of larvae were reared in mass culture under similar environmental conditions but in water filtered with 363 micron nitex screen cloth. Some specimens from one of these later hatches were preserved in 70% isopropyl alcohol. Larvae from other broods were not preserved. Details of those rearings, including data on growth of juveniles, are reported elsewhere (Brownell, Provenzano, and Martinez, in press). Measurements of zoeal carapace length (CL) were made from anterior- most margin of the eyestalk (prezoea, stage I) or rostrum (stage II) to the posterolateral carapace margin as seen in lateral view. Carapace length of the megalopa was taken from the tip of the rostrum to the tip of the projection on the posterior carapace margin, in lateral view. Carapace length of the first crab was measured from the notch between the rostral spines to the medial posterior carapace margin. Initially whole animals and molts were stained in acid fuchsin red, but the presence of flesh in whole animals, which obscured details, and the paucity of molts in good condition required utilization of additional whole specimens digested in hot 5% KOH and then stained in an aqueous solu- tion of chlorozol black. The resultant preparations were most satisfactory. Dissections were done in lactic acid and appendages were mounted in glycerine jelly. Whole drawings were made with the aid of a Wild M-8 stereomicroscope with camera lucida. Appendages were illustrated using a Unitron binocular VOLUME 90, NUMBER 3 737 compound microscope with attached camera lucida. All scales on figures are in mm. The 2 females which hatched the larvae studied in this paper and a male which copulated with one of the females subsequent to hatching of her brood have been deposited in the National Museum of Natural History, Smithsonian Institution. Results In the development of this species, as is usual for majid crabs, there are 2 zoeal stages and a megalopa. The first zoeal stage is preceded by a non- swimming prezoeal stage lasting not more than a few hours. Observations on 7 hatches from 4 different females indicated that at temperatures of approximately 24-28°C, time from hatching to megalopa is approximately 60-72 hours and from hatching to first crab stage approximately 130-148 hours. No meaningful information on larval survival rates was obtained. During some of the rearings no aeration was used, and in others aeration was irregular. Description of Developmental Stages Prezoea (Fig. 1) Duration.—Less than 12 hours. Size—CL 1.2-1.3 mm. In the earliest hatched (prezoeal) form, the dorsal carapace spine is not extended and the setae of the maxillipeds are not evaginated. Pereiopodal buds are already present, as are pleopod buds and spines on the abdominal somites (Figs. 1A, 1B). Carapace with blunt process representing dorsal carapace spine, rostral spine small, ventrally projecting. A forehead protuberance located dorsal to the ocular peduncles. Eyes sessile. Pereiopodal buds unarmed, first pair chelate (Fig. 1C). Abdomen consisting of 5 somites plus telson. Somite 2 with a lateral knob projecting slightly forward on each side, somites 3-5 with prominent posterolateral spines. Telson forked (Fig. 1D), but forks not fully extended, lateral spine on each side, 3 setae on either side of median notch. Antennule conical, with 2 long and one shorter terminal aesthetascs (Fig. 1E). Antennal protopod not fully extended in prezoea, an endopodal bud proximally, not reaching past tip of unextended protopod. Exopodite with a large terminal seta and 2 smaller subterminal setae (Fig. 1F). Mandible not studied. 738 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 1. Mithrax spinosissimus, prezoea. A, Dorsal view; B, Lateral view; C, Pereiopodal buds; D, Telson; E, Antennule; F, Antenna; G, Maxillule; H, Mazxilla; I, First maxilliped; J, Second maxilliped. VOLUME. 90, NUMBER 3 - af 739 Maxillule with unsegmented (?) endopodite, bearing 2 short setae (Fig. 1G) terminally. Basal lobe with 3 thick and 2 shorter, finer setae. Coxal lobe with 6 short, simple setae. Maxilla with well developed scaphognathite bearing 15 to 16 short setae marginally. Endopodite with 2 short simple setae, lobes of bifurcated basal endite each with 2 similar short setae, lobes of bifurcated coxal endite each with a single seta (Fig. 1H). First maxilliped with exopodite bearing 4 bifurcate short natatory setae resembling pairs. Five-segmented endopodite without setation except 3 short setae on terminal segment (Fig. 11). Second maxilliped with unsegmented endopodite bearing 2-3 very short simple setae, otherwise similar to first (Fig. 1J). Third maxilliped biramous, unsegmented, unarmed. First Zoea (Fig. 2) Duration.—1.5-2.0 days. Size.—CL 1.0 to 1.2 mm. Appendages and setation fully extended. Carapace with a moderately developed posteriorly hooked dorsal spine. Forehead protuberance present. Rostral spine small, projecting ventrally between antennules. Eyestalks fused to carapace. Pereiopodal buds larger, otherwise unchanged. Ab- domen with 5 somites plus telson. A lateral forwardly directed knob on each side of somite 2. Somites 3-4 with prominent posterolateral spine on each side (Figs. 2A, 2B). Telson sharply forked, each side bearing a lateral spine and 3 short setae medially, prominent medial notch (Fig. 2C). : Antennule conical, 3 long and 2 slightly shorter aesthetascs terminally (Fig. 2D). Antennal protopod extending beyond tip of exopodite. Exopodite with a strong terminal seta and 2 small ones subterminally. Endopodite shorter than exopodite, unsegmented, unarmed (Fig. 2E). Mandible simple, bearing many teeth. Maxillule with unsegmented endopodite bearing 2 very short, simple terminal setae. Basal endite with 6 setae, coxal endite with 5 simple setae (Fig. 2F). Maxilla with scaphognathite bearing 30 marginal setae, endopodite with 1 subterminal simple seta. Each lobe of basal endite also with 3 simple setae, lobes of coxal endite with only 1 each (Fig. 2G). First maxilliped with 4 natatory setae on exopodite, 5-segmented endop- odite bearing 3-4 setae on terminal segment, 2 distally on penultimate segment. One each on next 2 segments. No setae on proximal segment of endopodite (Fig. 2H). 740 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON VOLUME 90, NUMBER 3 741 Second maxilliped with 4 natatory setae on expodite, unsegmented en- dopodite in some bearing only 1 short simple seta subterminally, others bear 1 terminal, 2 subterminal at locations apparently suggestive of segmentation (Figs. 21, 2J). Third maxilliped present as bud, bilobed, with endopodite showing traces of segmentation, exopodite simple, unarmed. Second Zoea (Fig. 3) Duration.—1.0-2.0 days. Size.—CL 1.1-1.2 mm. The second stage differs little from the first, the major distinctions be- ing separation of the eyestalks from the carapace (Figs. 3A, 3B) and minor changes in some appendages, notably the maxillipeds and the antenna. Eyestalks with papilla on anterior margin. Rostrum now more anteriorly directed, otherwise carapace not much changed. Pereiopodal buds showing distinct segmentation, but still nonfunctional (Fig. 3C). Abdomen essentially unchanged except for slightly longer pleopodal buds, telson unchanged (res SID) Antennules essentially unchanged, still with 5 aesthetascs, 3 of which are longer than the remaining 2 (Fig. 3E). Antennal endopodite now segmented, reaching to base of terminal spine of the exopodite, and with single short, simple seta (Fig. 3F). Mandible unchanged. Maxillule with endopodite now showing 2 setae much longer than in previous stage, basal endite with 5 larger and 2 smaller, simple setae, coxal endite with 5 short simple setae (Fig. 3G). Maxilla with 31 marginal setae on scaphognathite. Endopodite, and each lobe of basal endite with two subterminal setae, lobes of coxal endite each with single short simple seta (Fig. 3H). First maxilliped now bearing 6 natatory setae on exopodite, 5-segmented endopodite with 3-4 setae on terminal segment, at least 1 on other segments except the most proximal (Fig. 31). Second maxilliped also with 6 (rarely 5) natatory setae, endopodite un- segmented, bearing 2 simple subterminal setae (Fig. 3). Third maxilliped biramous, neither exopodite nor endopodite with any setae, endopodite showing segmentation (Fig. 3K). < Fig. 2. Mithrax spinosissimus, first zoea. A, Lateral view; B, Ventral view; C, Telson; D, Antennule; E, Antenna; F, Maxillule; G, Maxilla; H, First maxilliped; I, J, Second maxillipeds of 2 different specimens. 742, PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON C Fig. 3. Mithrax spinosissimus, second zoea. A, Dorsal view; B, Lateral view; C, Pereiopodal buds; D, Telson; E, Antennule; F, Antenna; G, Maxillule; H, Maxilla; I, First maxilliped; J, Second maxilliped; K, Third maxilliped. VOLUME 90, NUMBER 3 : 743 Fig. 4. Mithrax spinosissimus, megalopa. A, Dorsal view; B, Lateral view; C—G, First to fifth pereiopods; H, Telson. Megalopa (Figs. 4, 5, 6) Duration.—3-4 days. Size —CL 1.2-1.5 mm. Carapace about 40% longer than wide in dorsal aspect, widest at bran- chial region, but not markedly so. Rostrum (central process) exceeding anterior projections (rostral lateral processes) of carapace (apparently not, in dorsal view) and deflected ventrad slightly less than 45° from plane of dorsal surface of carapace. Carapace moderately inflated dorsally, with 744 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON { \ OR Ps Vb, E Ai Vy \ vi wy, M ‘ y ) NW : y D \ \y \ : v ean A Fig. 5. Mithrax spinosissimus, pleopods of megalopa. Left to right, pleopods of second through sixth abdominal somites, respectively. 3 median tubercular projections (gastric, cardiac and posterior median processes) and 3 more on each side, the most anterior (anterior gastric lat- eral) being approximately in line with anteriormost (gastric) median tuber- cle. A prominent seta and several smaller ones on dorsal margin of carapace over eyestalk (Figs, 4A, 4B). Pereiopods now functional, bearing numerous setae (Figs. 4C-4G). Sixth abdominal somite now distinct, bearing pleopods. Pleopods biramous, functional, endopodite with minute hooks, exopodites of somites 2-6 bearing 9-10, 11, 10-11, 9 and 5-6 setae respectively (Fig. 5). Telson with rounded posterolateral corners, posterior telson margin slightly concave (Fig. 4H). Antennules biramous, peduncle 2-segmented. Each ramus 2-segmented, dorsal flagellum with 3 and 5 aesthetascs on distal and proximal segments respectively. Terminal seta on distal segment and apparently another on peduncle proximal to ventral flagellum with 2 terminal setae (Fig. 6A). Antennal peduncle with minute nephropore on a slight protuberance near base. Coxal segment apparently fused with basal segment, but line not distinguishable. “Basal” segment of peduncle with a large, ventrolateral VOLUME 90, NUMBER 3 745 Fig. 6. Mithrax spinosissimus, appendages of megalopa. A, Antennule; B, An- tenna; C, Mandible; D, Maxillule; E, Maxilla; F, First maxilliped; G, Second mavilliped; H, Third maxilliped. 746 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON knob and a smaller ventromedial knob distally, 2 additional peduncular segments distally. Flagellum 3-segmented, proximal segment not as long as wide, middle segment long, and terminal segment shorter but still about 4 times its width, each flagellar segment bearing up to 3 setae (Fig. 6B). Mandible with cup-shaped molar surface and 2-segmented palp bearing about 5 setae on distal segment (Fig. 6C). Maxillule with unarmed, unsegmented endopodite. Basal endite with 12-15 setae, coxal endite with 8 (Fig. 6D). Maxilla with scaphognathite bearing 33-37 marginal setae. Endopodite unsegmented, with sharp tip but otherwise unarmed. Basal lobes bearing 5-6 setae each, coxal lobes 3-5 (Fig. 6E). First maxilliped with medial margin of protopodite bearing total of 16 setae, reflexed epipodite laterally, bearing 4 long setae. Two-segmented exopodite with 4-6 plumose setae; unsegmented endopodite unarmed (Fig. 6F). Second maxilliped lacking epipodite (and setae on propodite?). Two- segmented exopodite with 4-6 plumose setae, the apparently 4-segmented endopodite showing indistinct fusion of 2 segments proximally (Fig. 6G). Third maxilliped with reflexed epipodite bearing 4-6 long setae, 4 shorter ones at base of 2-segmented exopodite which carries 4 plumose setae termi- nally. Endopodite strongly developed, proximal of 5 segments expanded medially, bearing approximately 12 setae and 4 teeth on medial margin. Next distal segments with additional, variable numbers of setae (Fig. 6H). First Crab (Fig. 7) Duration—No data. Size —CL 1.5 mm. Rostrum visible in ventral view only, depressed and directed posteriorly. A pair of well developed rostral spines reaching to base of antennal flagellum. A large postorbital or hepatic spine, showing signs of bifurca- tion. More ventrally in the same region, another but slightly smaller spine. Three spines along each side of dorsolateral carapace in mesobranchial region. Three more laterally along ventral margin. A prominent spine on each eyestalk, anteriorly. A dorsal process located medially, in line with postorbital spines (Fig. 7A). Antennule with 3-segmented dorsal flagellum, 2-segmented ventral fla- gellum, former with about 9 aesthetascs, including only 1 terminally, the latter with 3 simple setae on distal segment (Fig. 7B). Antennal flagellum now with additional segment, otherwise little changed, but peduncle now bearing spinose process on a proximal segment, with a single large spine more proximally (Fig. 7C). Mandible similar to that of preceding stage, with rounded molar surface, 2-segmented palp bearing 6 short stout setae (Fig. 7D). VOLUME 90, NUMBER 3 747 Fig. 7. Mithrax spinosissimus, first crab stage. A, Dorsal view; B, Antennule; C, Antenna; D, Mandible; E, Maxillule; F, Maxilla; G, First maxilliped; H, Second maxil- liped; I, Third maxilliped; J—L, Pereiopods 1-3 respectively. 748 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Maxillule with unsegmented endopodite now bearing 2 or 3 simple setae, basal and coxal endites bearing approximately 11 and 5-9 simple setae respectively (Fig. 7E). Maxilla with 31-34 marginal setae on scaphognathite, endopodite with a single long simple, subterminal seta, lobes of basal endite bearing 6 and 5 setae, lobes of coxal endite each with 2 or 3 simple setae (Fig. 7F). First maxilliped damaged, but apparently with reflexed distal segments of exopodite bearing 4 or 5 long setae, endopodite simple, unsegmented, unarmed except for 2 or 3 short setae. Basipodite with about 14-15 setae, mostly on medial margin, coxal lobe with 5 or 6 (Fig. 7G). Second maxilliped with 5 terminal setae on exopodite. Endopodite with 3 short segments distally, about 8 setae on distal segment, 2 or 3 on pen- ultimate segment, 1 on distal margin of next segment. Most proximal, fourth segment unarmed, at least 2 times longer than wide. A long lat- eral epipodite bearing about 8 fine setae (Fig. 7H). Third maxilliped with 4 setae terminally on exopodite, proximal segment with 5 marginal and 2 or 3 submarginal setae. Endopodite 5-segmented, each segment bearing setae, proximal segment with strong teeth on medial margin laterally (Fig. 71). Pereiopods setose. A dorsolateral tooth proximally on propodus of che- liped, a single tooth on ventral margin of dactyl of Ps, 2 teeth ventrally on dactyl of P;. Other pereiopods not available on specimen (Figs. 7J-7L). Discussion Although larvae are known for representatives of 45 genera of majid crabs, few of these are closely related to Mithrax spinosissimus. Kurata (1969) reviewed larval characters of Majidae and presented a diagram of assumed relationships. Unfortunately his paper is mainly in Japanese, and he was unaware of the then unavailable work of Yang (1967). The latter investigator, who also reviewed the systematics of the Majidae from the point of view of larvae, has published some of his findings (Yang, 1968, 1971, 1976), and other works are in preparation. It is not our purpose to attempt a comprehensive review of larval characters in the family. We will restrict our discussion to the relatively few species of Mithracinae for which larvae are now known. Within Mithrax, first zoeal stages of only 2 other species have been described, M. forceps (Lebour, 1944) and M. pleuracanthus (Yang, 1967). In addition, Yang reared larvae of M. sculptus and M. forceps and referred to them briefly in his discussions, but did not present full descriptions. Therefore we are limited in our ability to compare larvae of M. spinosis- simus with others in the genus, no second zoeal stages or megalopal stages having been described for other Mithrax. Descriptions of first stage larvae only are available for Micippia thalia VOLUME 90, NUMBER 3 749 (Kurata, 1969) and Tiarinia cornigera (Aikawa, 1937; Kurata, 1969), while first and second stage zoea, and the megalopa are known for Macrocoeloma diplacanthus, M. camptocerum (Yang, 1967), and Microphrys bicornutus (Hartnoll, 1964; Lebour, 1944; Yang, 1967). Description of the megalopa stage is also available for Micippia thalia (Kurata, 1969). Although the only majid for which direct development is known (Paranaxia serpulifera) is also a mithracine crab (see Yang, 1967:152), we do not have access to a description or to specimens of the megalopa and must delete it from further consideration. The first zoeae so far described for Mithracinae can all be distinguished from each other on the basis of one or more features with the exception of Mithrax forceps for which certain appendages and other characters are not yet described. Mithrax spinosissimus differs from all the others in having 5 rather than 3 aesthetascs on the antennule, in having an unsegmented, nearly unarmed endopodite on the maxilla (data not available for Micippia thalia and Mithrax forceps), as many as 30 marginal setae on the scaphognathite of the maxilla (data not available for Micippia thalia and Mithrax forceps). It may also differ from all but Tiarinia in having a dorsal seta on the telson forks. Tiarinia cornigera is unique in having 4 marginal seate on the scaphognathite of the maxilla (no data for Micippia). It shares only with Micippia the presence of a medial telson notch. Micippia differs from all the other species in having lateral knobs on the third as well as the second abdominal somite and in lacking the dorsal carapace spine, but having lat- eral carapace spines. The 2 species of Macrocoeloma differ from the other species in each having 10 marginal setae on the scaphognathite in addition to a posterior process, and in having 5 setae on the distal segment of the endopodite of the maxillule. M. diplacanthum is unique in having the distal segment of the endopodite of the first maxilliped with an expanded spine rather than the normal seta on the dorsal margin, and further differs from M. camptocerum in having a concave rather than straight telson fork, and a constricted rather than normally rounded forehead. Microphrys bicornutus differs from Mithrax pleuracanthus in having 6 setae on the distal segment of the maxillular endopodite rather than 7 typical of M. pleuracanthus. These distinguishing features are summarized below in the form of a key to the described first zoeae of the Mithracinae. Key to Described First Stage Larvae of Mithracinae 1. Lateral carapace spines present, no dorsal carapace spine; rostrum much longer than antennule; lateral knobs on second and third abdominal somites Micippia thalia 750 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON ~ No lateral carapace spines, a dorsal spine present; rostrum very short; lateral knobs only on second abdominal somite 2 2. Each telson fork with a dorsal spine, a large and a small lateral spine Tiarinia cornigera — Each telson fork with only 1 lateral spine (no further separation of Mithrax forceps possible) 3 3. Scaphognathite of maxilla with 10 marginal setae in addition to posterior process; 2-segmented endopodite of maxillule with 5 setae on distal segment, 1 on proximal Macrocoeloma — Scaphognathite of maxilla with more than 10 marginal setae 4. Dorsal margin of distal segment of first maxilliped with an expanded spine rather than a normal seta; telson fork slightly concave, fore- head constricted Macrocoeloma diplacanthum — Dorsal margin of distal segment of first maxilliped with a normal seta; telson fork straight; forehead normal, rounded M. camptocerum 5. Scaphognathite with 12 marginal setae in addition to posterior process; endopodite of maxillule 2-segmented; no dorsal seta on straight telson fork; 3 aesthetascs on antennule 6 — Scaphognathite with 30 marginal setae; endopodite of maxillule unsegmented, unarmed except for 2 short spinules; dorsal setae on slightly concave telson fork; 5 aesthetascs on antennule Mithrax spinosissimus 6. Distal segment of endopodite of maxillule with 7 setae, proximal segment with 1; rostral spine reaches midpoint of antennule; dorsal carapace spine stout at base, basal portion projecting anteriorly then sharply backwards Mithrax pleuracanthus — Distal segment of endopodite of maxillule with 6 setae, proximal segment with 1; rostral spine reaches beyond middle of antennule; dorsal carapace spine slender near base, gently curving posteriorly Microphrys bicornutus OU 4 Tiarinia along with Macrocoeloma has been placed sometimes in a sepa- rate subfamily, the Macrocoelominae, but Yang, following Garth (1958) con- sidered Tiarinia and Macrocoeloma to fall within the Mithracinae. Never- theless, he pointed out a number of primitive characters of Tiarinia. Among these are the presence in the first zoea of only 4 marginal setae on the scaphognathite of the maxilla (in addition to the posterior process), pres- ence of 7 rather than 6 or the more typical 5 spines on each side of the telson, a bifurcate endopodite on the maxilla, and lack of pereiopodal and antennal endopodal buds. Yang suggested some of these features might indicate more than 2 zoeal stages in Tiarinia. The position of this genus relative to other majids is not clear (Yang, 1967:152, 237-238). Micippia first stage larvae, while sharing with Tiarinia a median telson notch, differ from those of that genus as well as from all 6 of the other VOLUME 90, NUMBER 3 751 described larvae of Mithracinae in having no dorsal carapace spine, posses- sing lateral carapace spines, and in having each side of the third abdominal somite as well as the second with a lateral knob. Hence, it does not ap- pear to be close to Mithrax. Yang has earlier commented on the close similarity of Mithrax and Mi- crophrys larvae and the similarities of both with Macrocoeloma. Recogni- tion of larval features of generic significance, if any, must await additional descriptions. The information available for Mithracine megalopal stages is even more limited, for in addition to Mithrax spinosissimus, this stage is known only for Micippia thalia (Kurata, 1969), Microphrys bicornutus and the 2 species of Macrocoeloma (Yang, 1967). Few characters of significance have been described equally well for all species. There is some difference in size, as might be expected, but the megalopa of M. spinosissimus is approxi- mately the same size as that of Macrocoeloma diplacanthum, a species that reaches only a small size as an adult. The number of setae on the pleopods (uropods) of the sixth abdominal somite varies (Macrocoeloma camptocerum and Micippia thalia have 4, Macrocoeloma diplacanthum and Microphrys bicornutus have 5, Mithrax spinosissimus may have 5 or 6) and setation of other pleopods may vary with species. The setation of the scaphognathite, known for only 3 of the 5 species, may prove to be of some specific value. Yang (1967) tabulated features of 14 megalopal stages of majid crabs of known parentage. He established 7 groups of megalopal stages within the family, placing Microphrys, Macrocoeloma (Mithracinae), Pitho (Ophthalmi- inae), and Epialtus (Acanthonychinae) together for sharing small lateral ridges on the gastric region, and a small rostral spine bending downwards (except for Microphrys). Characters unique to the megalopal stages of Mithracinae or to genera within it are not yet known. Acknowledgments The experimental work was done while W.N.B. was with the Fundacion Cientifica Los Roques, Caracas, Venezuela. We are grateful to the mem- bers of the Fundacion and to its field staff for support in this work. The study of material and preparation of the manuscript was supported by Na- tional Science Foundation grant DEB 76-11716 to Old Dominion University Research Foundation. Mr. Joseph W. Goy prepared the illustrations. Dr. R. B. Manning kindly confirmed the identity of the adult crabs. Literature Cited Aikawa, H. 1937. Further notes on brachyuran larvae. Rec. Oceanogr. Works Japan 9(1):87-162, 36 figs. Brownell, W. D., A. J. Provenzano, and M. Martinez. In press. Culture of the 752 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON West Indian giant spider crab, Mithrax spinosissimus, at Los Roques, Venezuela. Proc. World Mariculture Soc., Eighth Ann. Meeting, Held at San Jose, Costa Rica. Louisiana State University, Baton Rouge, Louisiana. Garth, J. S. 1958. Brachyura of the Pacific coast of America. Oxyrhyncha. Allan Hancock Pacific Exped. 21(1-2):1-854, pls. A-Z, 1-55. Hartnoll. R. G. 1964. The zoeal stages of the spider crab Microphrys bicornutus (Latr.). Ann. Mag. Nat. Hist., ser. 13, 7:241-246, 3 figs. Hazlett. B. A.. and D. Rittschof. 1975. Daily movements and home range in Mithrax spinosissimus (Majidae, Decapoda). Mar. Behav. Physiol. 3:101-118. Kurata, H. 1969. Larvae of Decapoda Brachyura of Arasaki, Sagami Bay-IV. Majidae. Bull. Tokai Reg. Fish. Res. Lab. No. 57:81—-127, 27 figs. Lebour, M. V. 1944. Larval crabs from Bermuda. Zoologica, N.Y. 29(3):113-128, 19 figs. Williams, A. B. 1965. Marine decapod crustaceans of the Carolinas. Fish. Bull. U.S. Fish. Wildl. Serv. 65(1):1-298, 252 figs. 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, 1459, 94 figs. —. 1968. The zoeae, megalopa and first crab of Epialtus dilatatus (Brachyura, Majidae) reared in the laboratory. Crustaceana, Suppl. 2, Studies on decapod larval development: 181-202, pls. v, vi, 9 text-figs. 1971. The larval and postlarval development of Parthenope serrata reared in the laboratory and the systematic position of the Parthenopinae (Crustacea. Brachyura). Biol. Bull., Woods Hole 140(1):166—189. ——. 1976. Studies on the western Atlantic arrow crab genus Stenorhynchus (Decapoda, Brachyura majidae). I. Larval characteristics of two species and comparison with other larvae of Inachinae. Crustaceana 31(2):157—-177. i - arg i * erst : FR i ; : is apa oa i i Oia ea pole a i oss < i pier a= " is 5 _ - a ere ‘ aoe ” ae oan) : t i pu ite i 7 ? i hoy ee f t a ; ‘ i ? ~ ‘ a) i ; a f in { r : 1 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, Washing- ton, D.C. 20560. 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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 81% 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 indica- tions 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 retumed proof. CONTENTS The Soricidae of Taiwan. E. W. Jameson, Jr. and Gwilym S. Jones Three new species of Oligochaeta (Naididae) from the southeastern U.S. Walter J. Harman A new entocytherid ostracod of the genus Plectocythere. Arnold W. Norden Rediscovery of Sesarma aequatoriale Ortmann, 1894 in the eastern Pacific (Crus- tacea, Decapoda, Grapsidae). Lawrence G. Abele Studies on decapod Crustacea from the Indian River region of Florida. VI. The identity of Parthenope (Platylambrus) serrata (H. Milne Edwards, 1834) — and Parthenope (Platylambrus) granulata (Kingsley, 1879). Robert H. Gore The larva of a new subterranean water beetle, Haideoporus texanus (Coleoptera: Dytiscidae: Hydroporinae). Glenn Longley and Paul J. Spangler The Sphegina species of eastern North America (Diptera: Syrphidae). Gary A. Coovert and F. Christian Thompson Key to the Neotropical genera of Parydrinae with a revision of the genus Eleleides Cresson (Diptera: Ephydridae). Wayne N. Mathis Three new Ecuadorian species of the aquatic beetle genus Chaetarthria (Coleop- tera: Hydrophilidae). Paul J. Spangler Two new species of Colobomatus (Copepoda: Cyclopoida) parasitic on Pacific fish. Roger F. Cressey Four new species of Anoplodactylus (Pycnogonida) from the western North At- lantic C. Allan Child Range extensions of the shrimps Solenocera necopina Burkenroad and Parapenaeus americanus Rathbun (Crustacea, Decapoda, Penaeoidea). Isabel Pérez Farfante New entocytherid ostracods of the genus Dactylocythere. Horton H. Hobbs, Jr. and Margaret Walton Natural history of Plummers Island, Maryland. XXII. Biting midges (Diptera: Ceratopogonidae). 1. Introduction and key to genera. Willis W. Wirth, Niphan C. Ratanaworabhan and Donald H. Messersmith Description of a new hermit crab (Family Paguridae) from southern California and Mexico. Janet Haig Orchestia vaggala, a new land-hopper from the Galapagos Islands (Crustacea: Amphipoda: Talitridae). Thomas E. Bowman A new frog of the genus Rana from the Philippines. Walter C. Brown and Angel C. Alcala Sarsiella maurae, a new species of marine Ostracoda (Sarsiellidae: Myodocopina) from Bahia de Los Angeles, Gulf of California, Mexico. Louis S. Kornicker Taxonomic status of the turtle, Chrysemys picta, in the northern peninsula of Michigan. Carl H. Ernst and James A. Fowler A great auk, Pinguinis, from the Pliocene of North Carolina (Aves: Alcidae). Storrs L. Olson Natural history of Plummers Island, Maryland. XXIII. Studies on lichen growth rate at Plummers Island, Maryland. James T. Lawrey and Mason E. Hale, Jr. A gill-inhabiting new genus and species of the Branchiobdellida (Annelida: Clitellata). Perry C. Holt Larval and early post-larval stages of the West Indian spider crab, Mithrax spinosissimus (Lamarck) (Decapoda: Majidae). Anthony J. Provenzano, Jr. and W. N. Brownell 459 483 491 495 505 532 536 553 566 579 584 597 600 615 648 658 669 676 685 690 698 726 735 4 577,0673 Proceedings of the BIOLOGICAL SOCIETY of WASHINGTON Volume 90 6 December 1977 Number 4 THE BIOLOGICAL SOCIETY OF WASHINGTON 1977-1978 Officers President: Richard S. Cowan Secretary: W. Duane Hope Vice President: Clyde F. E. Roper Treasurer: Oliver S. Flint, Jr. Custodian of Publications: John H. Miles Elected Council J. Laurens Barnard Raymond B. Manning Ronald I. Crombie John H. Miles Frank D. Ferrari PROCEEDINGS Editor: Austin B. Williams Coeditor: C. W. Hart, Jr. Associate Editors Classical Languages: George C. Steyskal Invertebrates: Thomas E. Bowman — Plants: David B. Lellinger Vertebrates: Leslie W. Knapp 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 Institution, Washington, D.C. 20560. The Proceedings of the Biological Society of Washington is issued quarterly. Manuscripts, corrected proofs, editorial questions should be sent to the Editor, Biologi- cal Society of Washington, National Museum of Natural History, Smithsonian Institution, Washington, 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 Application to mail at second class postage rates is pending at Washington, D.C., and additional mailing office. PROC. BIOL. SOC. WASH. 90(4), 1977, pp. 753-763 OBSERVATIONS ON SPECIMEN FIXATION William R. Taylor This study was initiated to learn something of the possible value of glutaraldehyde as a fixative in comparison with formaldehyde, and to de- termine the value of buffered and unbuffered fixatives as long term pre- servatives for whole specimens of fishes relative to changes in transparency (clearing) and color, resistance to breakdown and clearing by pancreatin, and ability of ossified tissues to stain after one year in preservation. Although Miller (1952) recommended the initial addition of one teaspoon of sodium borate (borax) to a half gallon of 10% formalin (approximately 1.1 g/900 ml) and Taylor (1967, p. 6) suggested the addition of a similar quantity after a day of fixation, the present study indicates that borax should be avoided. Furthermore it may not be desirable to buffer formal- dehyde solutions to alkaline pH because increase of pH alkalinity above 7.0 seems to enhance autolysis as evidenced by clearing of specimens. Instead, a solution of fixative saturated with powdered limestone (or possibly marble chips) may be a better preservative. I wish to thank the following for aid in various ways in this study: Martin L. Wiley for aid in collecting specimens; Eugene Jarosewich, Leslie W. Knapp and John C. Harshbarger for use of equipment and supplies; and George Van Dyke for laboratory aid. I am grateful to Robert H. Gibbs, Jr.. Ernest A. Lachner and Stanley H. Weitzman for comments on the manuscript. Materials The following materials were used in the preparation of the 12 fixatives: Glutaraldehyde; 50% solution of glutaraldehyde, biological grade, pH. 3.2. Formalin; approximately 38% solution of formaldehyde, USP, pH 3.4. Borax; sodium borate powder, USP. Distilled water (single simple distilled); approximate pH 6.3. Limestone; powdered, commercial grade. Saturated borax solution; the clear supernatant from a saturated solution of sodium borate in distilled water, pH 9.2. Limestone water; a solution, approximately 3.3% saturated (prepared from the clear supernatant of a saturated solu- tion of limestone in distilled water, saturated pH 9.0), pH 7.2. List of Fixatives 1. Formalin, 10% (approximately 3.8% formaldehyde), in distilled water. 2. Same as number 1, but added 1.1 g borax per 900 ml at 19 days. 3. Formalin, 10%, in saturated borax solution. 754 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 4. Same as number 3 for 18 days; drained off solution on 19th day, rinsed and changed to 10% formalin in distilled water. Same as number 1, but with 1.1 g borax per 900 ml. Formalin, 10%, in limestone water plus 1.1 g borax per 900 ml. Same as number 6, but lacking borax. Same as number 1, but with about three teaspoons limestone per 900 ml, forming permanent precipitate at bottom of container. 9. Formalin, 10%, plus 4% glutaraldehyde (pH of combination 3.2), in distilled water. 10. Same as number 1 initially; drained off solution on 7th day, rinsed and changed to 4% glutaraldehyde in distilled water. 11. Glutaraldehyde, 4%, in distilled water. 12. Glutaraldehyde, 4%, in saturated borax solution. Ce SPN Procedure Glass quart jars, sealed with rubber gaskets, were used. Since each jar held only slightly more than 900 ml, all solutions including specimens were adjusted to 900 ml. This left a minimum of free air space above the preservative. The specimens were considered as water in the total volume of solution in each jar, occupying about 4% and not more than 5% of that total. The specimens therefore tended to dilute the buffering solu- tions used, but did not materially increase the concentration percentages of the fixatives. Thus in fixative number 3, approximately 5% of the volume was considered water from specimens, about 3.8% formaldehyde, 6.2% water and other chemicals from the formaldehyde solution and 85% saturated borax solution. In fixative number 9 the total volume included about 5% specimens, 3.8% formaldehyde, 6.2% water and other chemicals from the formaldehyde solution, 4% glutaraldehyde, 4% water and other chemicals associated with the glutaraldehyde solution, and 77% distilled water. The basic materials used in this study were prepared in advance, but the combinations, listed for each fixative, were not made until specimens were added in the field. The fish specimens used were collected at the same time and place near Solomons, Maryland, 26 October 1974. A total of 18 specimens was fixed in each jar, including two species of Atherinidae, two cyprinodontids, one poeciliid and one centrarchid, except in some of the glutaraldehyde solutions in which a sciaenid was fixed instead of the centrarchid. Each species was distributed in equal numbers to each jar except in the case of the centrarchid. Specimens of as nearly equal size as possible were selected to provide uniformity of interpretation of results. Because of lack of pH meter in the field, the first pH readings were made about 24 hours after initial specimen fixation. These readings were made daily thereafter for a one-week period, and at subsequent times throughout one year. se ee ee ene ne a VOLUME 90, NUMBER 4 759 At the end of one year, two specimens from each lot were cleared with pancreatin and stained with alizarin Red S by the method of Taylor (1967) to observe loss of calcium minerals and resistance to enzymatic breakdown. Two additional specimens were stained but not cleared. All solutions used to compare differences in fixation were prepared in equal volumes and mixed thoroughly to provide standard conditions for interpretation of results. Observed Results Changes in pH.—The first week’s shift in pH in fixatives with speci- mens was generally toward neutrality in unbuffered solutions, but those buffered with borax or a combination of borax with limestone water showed little change or a very slight increase in alkalinity. Borax combined with glutaraldehyde formed a yellowish emulsion and precipitate with increase in acidity. After one year the observed pH was decidedly more acidic in all cases except two, solutions in which the fixative was saturated with powdered limestone (8) and another in which saturated borax solution was added to formalin (3). Powdered limestone acts as a weak buffer in formaldehyde solutions in contrast to sodium borate, a strong buffer. The approximate maximum pH readings for saturated solutions of borax followed by limestone are: in distilled water 9.2, 9.6; in 10% formalin in distilled water 9.1, 8.4. Addition of specimens will cause the pH to change almost instantly to 8.4-8.8 in borax- formalin solutions and to as acid as pH 5.6 with specimens crowded in a limestone-formalin solution. The change in pH is a result of the reaction of the specimens with formaldehyde. In contrast to alkaline buffered forma- lin solutions, specimens placed in unbuffered formalin (pH 3.7-4.2) produce a slight initial change in pH to about 5.6. The change in pH depends upon the volume of specimens added; few specimens produce the least change, whereas crowding specimens into a container produces the greatest pH alteration in both buffered and unbuffered solutions. The reaction of formaldehyde with proteins results in a layered effect in a container. Unless the solution is mixed frequently during the first few days of fixation the upper liquid may be one or two full pH units different from the liquid surrounding specimens. For example, in a limestone-formalin solution the pH may be 8.4 in the upper liquid and 6.4 at the specimen. The differential has been demonstrated by me to persist for days (and _ per- haps will continue for weeks) if the contents of the container are not dis- turbed. Consequently, the upper liquid in unbuffered solutions is more acidic than that near specimens whereas the liquid in basic buffered solu- tions is more acidic near specimens, the upper liquid remaining more alkaline. Following initial fixation there is a gradual slight rise toward alkaline pH in unbuffered solutions. 756 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 1. Specimens of Menidia beryllina illustrating translucency caused by retention in a formalin plus borax solution at alkaline pH, contrasted with opaque specimens re- tained in an unbuffered formalin solution. All specimens were fixed at the same time. The specimens were photographed in distilled water over a light box, about six years after collection, by George Van Dyke and the Smithsonian Photographic Services. From these observations, it is apparent that formaldehyde solutions undergo three shifts in pH in the presence of specimens: (1) An initial rapid increase in acidity by reaction of the formaldehyde with proteins (masked in acid solutions), followed (2) by a slight increase in alkalinity primarily resulting from the alkaline body fluids, and (3) a slow increase in acidity, presumably due to production of formic acid. Translucency.—lI have known for some time that specimens may clear VOLUME 90, NUMBER 4 757 in some formaldehyde solutions. The degree of clearing may be slight or extensive (Fig. 1). In this study I have rated the degree of clearing by comparison of specimens, side by side over a light box as: extensive— specimens translucent; considerable—specimens showing considerable trans- lucency in thinner areas of body; and slight or very slight—where little or no translucency exists. A numerical rating from 1 (most translucent) to 12 (virtually opaque) was also performed and the results are presented in Table 1 to express differences between those listed in the categories of considerable trans- lucency and very slight translucency. All specimens fixed in borax buffered solutions where the pH remained at or more alkaline than 7.0 for a lengthy period (2, 3, 4, 5, 6, 12) exhibited the greatest transparency. The specimens in formalin buffered with a saturated borax solution (3) were sufficiently cleared that objects placed between them and a strong light could be observed through them as shadows. The specimens in fixative number 2, initially fixed in acid formalin followed by fixation in formalin and borax, exhibited greater clearing than did those in unbuffered solutions (1, 9, 10, 11) or in solutions buffered with limestone water (7, 8). The formalin plus saturated limestone (8) and the unbuffered glutaraldehyde solutions (10, 11) exhibited the least clearing. An example (not part of the study reported here) of translucency re- sulting from the use of borax in a formalin solution is shown in Fig. 1. The specimens of Menidia beryllina illustrated were all collected and fixed at the beach, Chesapeake Biological Laboratory, Solomons, Mary- land, 3 June 1970 by Martin L. Wiley. They were placed in two screw-top jars containing similar formaldehyde and distilled water solutions. The jars were approximately two-thirds filled after addition of specimens. An un- known quantity of borax was added initially to one jar. On 9 September 1971 (463 days after collection) I noted that the liquid in one jar was yellowish and the specimens cleared, while in the clear whitish liquid of the other jar, the specimens were opaque. The pH readings that day were 8.1 for the former and 5.7 for the latter. There was no precipitate of borax in either lot at that time, but it was obvious (neither lot was labeled as buffered) that the borax had been added to the lot with the cleared specimens because of a pH of 8.1. On 27 February 1976 I obtained pH readings of 7.5 and 5.1, respectively for the same two lots. Color—All specimens retained in formalin solutions are whitish with dark pigment patterns evident, but those in borax buffered formalin gradually assumed a darkened cast. The specimens fixed in basic glutaral- dehyde (12) became dark brown with obscured pigment. Those in acidic glutaraldehyde (11) were dark yellowish brown with obscurement of pig- mentation. The specimens fixed in acidic formaldehyde followed by Z O a S) Z — cn nN << 0 es< aicie poinosqo yuoursid = ‘UMOIG Yep ALOA a], qeioprisuoo G 61> €'Q> 229599 ZI 5 Grace = eo= = cl pernosqo Ayjensed yuoursid va Te ae SUBLOIE| AKO] OS 1UG| qwysys ATQA TT) EG 8°g Sg € od SEG [EAS] PHISIEROD) V CL> o8> Sas Gte ©) = SQ ee od afqeiepisuoo ZL 68 Gg—sg ¢ VI-L G> V Pe] 6LOLE = pouoylep ynq ¢# Ur sy a] qetopIsuoo © 0'L> ‘a°9 08? EL Ie) GOSSto nc © SO-CL je Q younstp yuowsid iS SYSIIYA ‘AIOATIS WYSys AOA 6 peo 19 rg, added on each side. foretarsus of a form from Mt. Mitchell is presented as Fig. 2, A-B. The sensillum b’-1 occupies a peculiar postion in that it is distal to the seta delta 4’ and not proximal to it as in other Eosentomon species. The fe- male genital apparatus of the same specimen from Mt. Mitchell is shown as Fig. 2, C. On the holotype this structure cannot be seen clearly. Posi- tions of the setae and sensillae on terga VII-VIII are illustrated by Fig. 2, D. They are the same on the holotype. Glands of filamento de sostegno in prothorax. Ratios averaged from approximately 100 individuals: LR 3.1; ise aeivon 0:43: FU 93; TR 63; TDP 0:70. Discussion In the 114 adults, 8 larvae I, 5 larvae II, and 16 maturus junior there were no variations in: (a) presence of very long labrum and stylet-shaped mandibles; (b) presence of sensillum b’-1 on the foretarsus; (c) absence of club on pretarsal sensillum s; (d) setae (sensillae) P 1’ on tergum VII short, peglike, and inserted along the posterior margin of the sclerite; and (e) glands of the filamento de sostegno in the prothorax as they are in the type of E. rostratum. The chaetotaxy for each stage and the setae added in each are shown in Table 1. Prelarva and preimago stages not found. Twenty-six of the adults had discrepancies in setae, but every case was an obvious asymmetry because it involved the addition or loss of a single setae from only one side of the segment. The holotype also has nine setae in the anterior row on tergum VI instead of the usual eight. Several body structures and ratios used in the descriptions of Protura were measured and treated statistically to check their variation in this population. In some specimens the limits of certain of these structures 784 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON LR-Head rs TR-Tarsus A ee : aa II Bees ut | —_— 0.2 0.3 0.4 0.5 0.6 4.0 0) 6.0 7.0 EU-Tarsus TDP-Tarsus . arenes | iz m | eer | - ae I ee. 4 I rere I Sans Ogee O26S 0.9 0295-120 0.6 OST 0.8 0.9 Fig. 3. Comparison of the six ratios PR, LR, TS, TR, EU and TDP among four stages of Eosentomon rostratum. A, Adult; M, Maturus junior; II, Larva II; I, Larva I. could not be determined, and these were not included in the calculations. The median, standard deviation from the mean and standard error of the mean were calculated on the basis of the number of characters measured. The results are shown in Table 2. Lengths and ratios of the head and VOLUME 90, NUMBER 4 ~l CO Ol si S ¥0 A 40 30 30 20 20 10 WE) // 12 13 /% 15 50 40 30 20 1/0 0.37 04% O43 O46 O49 OF2 IP EI (RO AGA G1) 50 50 $O $0 3jO 30 rae) 20 /O /O O08) Of4 O87 090 093 09% 0.99 0.64 067 O7 O73 0.76 Fig. 4. Frequency distribution of six ratios tor adult Eosentomon_ rostratum. A, Pseudocular ratio (PR); B, Labral ratio (LR); C, Rostral setae 3:1 ratio (RS); D, Tarsal ratio (TR); E, Empodium-unguis ratio (EU); F, Total distal/proximal ratio (TDP). tarsus appear to be less variable than lengths and ratios of setae and sensillae. The more variable nature of seta and sensillae may be more apparent than real because errors in measurement of the smaller struc- tures has greater influence on the results obtained. The data indicate that of the foretarsal sensillae, t-1, e, g, and f-l1 show the least variation in length, but this may also be due to the greater difficulty in seeing and accurately measuring the others. 786 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Table 2. Variation in lengths and ratios of certain structures of adult Eosentomon rostratum Ewing. Std. Std. Structure* No. Min. Max. Range Median dev. error Head lengths Capsule H 107 150. 169.2 19.2 156.8 4.1 0.395 Labrum JL 46.8 56.4 9.6 50.9 1.78 0.168 Ratio eRe On 2.85 3.34 0.49 3:10) OOO OLO Total TL 107 198. PN. 24. DOK 4.06 0.488 Pseudoculus IP 89 10.8 14.4 3.6 1192) 01620065 Ratio PR 88 11.4 WD O20 13.18 0.78 0.083 Rostral setae RS Teel O2 18. 30. IP, 22'6Sin les AON? RS TL) 48. 70.8 22.8 52.12 ies S Ratio RS III/I 104 0.36 0.53 (O37 0.43 0.03 0.003 Tarsal lengths Empodium E 106 14.56 20.8 6.24 18.89 1.01 0.098 Unguis WO" Se? 22.36 4.16 20.42 0.82 0.079 Ratio HU elLOG 0.80 0.98 0.18 0.93 0.04 0.004 Distal . TD 108 48. 58.8 10.8 53.30) Some Onk9 Proximal 1 70.8 81.6 10.8 (G0 wae Oss2 Ratio TD 108 0.63 0.75 0.12 0:70 O03 01003 Total EL. 108 122.4 138. 15.6 128.77 3:0 OLS 36 Ratio TR 104 Seal 6.92 WL PAll 6.35 0.26 0.026 Mesonotum Setae pelt 107 33.6 44.4 10.8 3.02) 2204 Ono 7 jo) 1s 34.8 44,4 9.6 37.8 Deal OB al 74 18. 26.4 8.4 2214 ~~ ASG OIG au Wl 36. 48. 124, 41.9 262, Orsial: Abd. tergum IV Setae p l 109 32.4 42. 9.61 36.44 1.98 0.190 jo) Jo 27.6 36. 8.4 328 2 Ole On98 Tarsal sensilla tal: 109 7.8 11.96 4.16 9.66) (OMA 0070 te 109 He, 19.76 6.76 16.28 1.48 0.141 tes 24 D4 9.88 4.68 7.02; 7 el SaeeOL 36 ae 109 15.6 24.44 8.84 20.89 1.53 0.147 ley a 19 7.81 17.68 9.88 14.04 2.77 0.634 lo 102 10.92 20.8 9.88 15.6 2.01 0.199 Cc’ 18 ris) 18.2 10.4 12.48 254 0:780 XxX 109 P4016) 37.44 9.88 33.88 1.97 0.189 Y. 108 28.08 38.48 10.4 33.54 LAG 0.161 iL, 105 BL 40.56 9.36 35.94 77 Oss a 29 10.4 16.64 6.24 - 13.03 1.87 0.348 b 109 15.6 24.44 8.84 19.88 144 0.137 () 42 7.8 18.72 10.92 iy 3.09 0.420 oe ra ~l VOLUME 90, NUMBER 4 78 Table 2. Continued. Std. Std. Structure* No. Min. Max. Range Median dev. error d 45 6.24 16.12 9.88 Salle sei OAD € 104 11.44 17.68 6.24 13.44 0.95 0.093 g 109 11.96 16.64 4.68 TAO 0 0.096 dL 99 7.8 33, 5.2 10.11 0.96 0.097 f 2 22 7.28 11.44 4.16 10.2 VO5y 0225 s 101 14.56 22.36 7.8 IES) Oa 1! 0.139 Distance a’-t 1 107 Wy, 18.2 52, 15.63 0.94 0.091 Distance a’—t 2 106 22.88 29.64 6.76 22 lel Gs Ooleles Distance t 1-t 2 107 7.28 11.44 4.16 LO OA OOre * Lengths in microns. The ratios PR, LR, RS, TR, EU, and TDP for the four stages are com- pared in Fig. 3 and their frequency distributions in Fig. 4. Because of the small sample size, it is felt that only trends are indicated for each of the three immature stages. Literature Cited Berlese, A. 1909. Monografia dei Myrientomata. Redia 6:1-182. Bonet, F., and S. L. Tuxen. 1960. Re-examination of species of Protura described by H. E. Ewing. Proc. U.S. Nat. Mus. 112:265-305. Condé, B. 1961. Nouvelles récoltes de Protoures au Maroc. Bull. Mus. Hist. Nat. 33(5):495—499. Durey, R. A. 1963. A statistical treatment of Eosentomon pseudorostratum n. sp. M.S. thesis, East Tennessee State University, Johnson City, Tennessee. , and T. P. Copeland. 1968. Eosentomon pseudorostratum n. sp. (Insecta, Pro- tura, Eosentomidae) from North Carolina. Jour. Tenn. Acad. Sci. 43(2):60-63. Ewing, H. E. 1940. The Protura of North America. Ann. Ent. Soc. Amer. 33: 495-551. Imadaté, Gentaro. 1974. Protura-Insecta-Fauna Japonica. Keigaku Pub. Co., Ltd. 301 pp. Tuxen, S. L. 1949. Uber den Lebenszyklus und die postembryonale Entwicklung zweier danischer Proturengattungen. Kgl. Vid. Selsk. Biol. Skr. 6(3):1—50. —. 1964. The Protura. A revision of the species of the world. Hermann, Paris. 360 pp. Womersley, H. 1927. A study of the larval forms of certain species of Protura. Ent. Mon. Mag. 63:149-154. PROC. BIOL. SOGsAWASEH: 90(4), 1977, pp. 788-797 MYODOCOPID OSTRACODA OF THE INDIAN RIVER COMPLEX, FLORIDA Louis S. Kornicker Abstract —Kornicker, L. $., Department of Invertebrate Zoology, Smith- sonian Institution, Washington, D.C. 20560.—Five species of Ostracoda in the suborder Myodocopina are reported from the Indian River complex (Indian and Banana rivers), Florida; three species are reported from the continental shelf off east central Florida. Ostracode abundance increased during February to April, 1975, in the Banana River, and during February to May, 1975, in the Indian River. Diversity is low in the Indian River complex compared to that on the continental shelf. Reproduction is year- round for the two most abundant species in the complex. The Harbor Branch Consortium, Fort Pierce, Florida, has been con- ducting since September 1973, an investigation entitled the Indian River Coastal Zone Study. The study is supported jointly by the Harbor Branch Foundation, Inc., and the Smithsonian Institution. According to Dr. David K. Young, Principal Investigator for the project, one of the major goals of the investigation is to obtain baseline information on the diversity of organisms and quality of the environment in the Indian River region (Young, 1976:14). I have participated in the investigation by identifying the collections of Ostracoda in the suborder Myodocopina. The Indian River complex is a series of shallow lagoons extending about 160 miles along the eastern coast of central Florida. The com- plex includes the Indian River, Banana River, and Mosquito Lagoon. I received samples for study from the Indian and Banana rivers. In ad- dition, a few samples were collected on the continental shelf off east central Florida. Both the Indian and Banana rivers are separated from the Atlantic Ocean by a series of barrier islands breached by narrow inlets at St. Lucie, Fort Pierce, and Sebastian (Fig. 1). Freshwater runoff into the lagoons (mostly from May through October) combines with tidal ex- change through the inlets to impart an estuarine character to the la- goons (Eilers, 1975:4). Tidal influence, however, is felt mostly in the southern half of Indian River in the vicinity of the inlets. The northern part of Indian River and the whole of the Banana River are minimally af- fected by oceanic exchange through the inlets (Young, 1976:15). Addi- tional information concerning the physical, chemical and biological en- vironment of the Indian River complex and of the continental shelf off Florida are included in annual reports of the investigation for 1973- VOLUME 90, NUMBER 4 789 28° Ft. Pierce ~\ St. hucies Zt Fig. 1. Indian River complex and adjacent continental shelf. 1974, 1974-1975, and 1975-1976 (Young et al., 1974; Young [ed.], 1975, 1976). The size frequency of sediment in the sample area is given in Tablet. The samples from the Indian and Banana rivers containing ostracodes 790 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Table 1. Size frequency of sediment (mean weight percent) in vicinity of stations where ostracodes were collected. Analyses by R. T. Wolcott. Sand fraction (phi —1-4) determined by Ro-Tap sieve; Silt (phi >4->8) and clay (phi >9->10) by pipette analysis. Grain Haulover Banana Sebastian Link Port size Canal River Inlet 8 May 1975 phi 8 Aug. 1975 27 Oct 1975 27 Oct. 1975 5 Aug. 1975 al 0.7 0.2 1.9 0.5 0 DB 0.2 1.3 0.3 il 3.8 9.5 46 Shi 2 8.4 47.8 16.8 56.4 3 59.4 Stal OS 22.0 4 13.4 3.0 40.6 Sell >4 2.0 0.5 ILS 2.4 5) 2.0 0.4 13 DA) SE 1.8 0.4 133 ae, Sr 1.8 OF IL 1.8 >8 1.6 0.3 1.0 seal >9 1.4 0.2 0.8 1.4 >10 iL-33 0.1 0.6 1.0 were collected using a “box core” sampler which removed a sediment plug 225 cm? in area and 20 cm in depth (for illustration see Young, 1976: 15). The sampler is limited to use in water depths up to 3 m. Up to 5 samples were obtained at each station and some, but not all, stations were sampled monthly. In the field, all samples were sieved through 0.5 mm or 1.0 mm mesh nylon (“Nitex”) screening, narcotized (0.15% propylene phenoxy- tol in sea water), stained (rose bengal) and fixed (5-10% formalin in sea water). After 24-72 hours, samples were washed with fresh water and stored in 70% ethyl alcohol. Animals were sorted from sample residue us- ing dissecting microscopes. I also collected a non-quantitative sample using a smal] aquarium net in the Indian River near Vero Beach in October 1971. Samples from the continental shelf off Fort Pierce were collected using a Smith MacIntyre Grab by personnel aboard the R/V Gosnold, during February-March 1974. Detailed station data will be presented in a paper in preparation concerning the distribution of ostracodes in the western Atlantic and Gulf of Mexico. Geographic Distribution The “box core” sampler which removed a sediment plug of 225 cm? gen- erally contained only a few specimens of myodocopid ostracodes, but a suf- ficient number of samples were collected in the study area to give a general picture of ostracode distribution in the shallow areas sampled. 191 VOLUME 90, NUMBER 4 5888S ee 0 0) 0) I I GE 6 GV G (M.u8T.OT08 “NuPS.OToLZ) PIUL PONT 4S TON 0 0 0 I 0 9€ IG SI I (MuPSLT 008 “Nu8Sl9GoLZ) PUI 240d HOY 1PON 0 I 0 g 0) €E—-0G €c-61 ro am Og SF (MuPS.0G08 “Nu90/GEoLZ) HOd AUT FPON 0 I 0 I 0 G'6G—LG G'0G-06 00G G (M.96.06008 “NuGLZEoLZ) [UL 401d HOA IPON 0 0 I 0 0 d d RO I (M.£6.008 “N.6EoLZ) YRC C19A 1VON 0 I 0 i I VE-GG CS-FI cy—-OF € (Mu9E/LE-08 “NuOETS.L3) [UL UeHseqes 1vON 0 0 0 O08 99 GV—GG Ve-91 OOI—0 GE (M.0E-SF008 “N.00.PP.8G) [RUBD TeAoineyY 1vON IOATY UPIpUy T 0 0 GG 0 COV-8T SCas 16 OF-EG €1 (M.i00.LE008 “N.00/G1-8G) TAT PuRUuBY ‘Jopul 8148112 DUD Dj0I sypapd 20/ ays asueI S9pPOdP.1}SO UOI}B00'T ‘ds -n90_--Xa}-— -1aySOZ = -SIP osuel osuel uo YIM adolajs Uu0laz Dyas DyJaIs ~~ Dians AyUTTeS ain} yidoq sojdures -DIDg -do1aysy -1DS -IDS = -ADS -elodula [, jo 1aquinNy nnn nnn ee EEE SSEES ESSE SESE ‘peyeTfoo alam Ady} 1oYM suoT}e}S JO Ayturyes pue ‘ginjzeroduis} ‘yydep 10zyeM Jo sasuei pue soloeds yore jo suoutloads jo sioquinNy “J 2a[qeL 792, PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON With the exception of a single juvenile of Parasterope sp. indet., the Banana River station produced only Sarsiella zostericola Cushman, 1906. That species and Sarsiella disparalis Darby, 1965 are common species in the Indian River (Table 2). No other species of Sarsiella was collected in the Harbor Branch sampling program, but a single juvenile of Sarsiella texana Kornicker and Wise, 1958, was collected by the author in the In- dian River near Vero Beach. Three specimens of Asteropteron oculitristis Darby, 1965, were collected in the vicinity of inlets in the Indian River and could be immigrants from the Atlantic. Three samples from the continental shelf off Fort Pierce from depths of 10-28 m contained 3 species, each represented by 1 specimen: Pseudo- philomedes ferulanus Koricker, 1958, Actinoseta chelisparsa Kornicker, 1958, and Cycloleberis americana (Miller, 1890). Seasonal Distribution Samples were collected more or less monthly from June 1974 through July 1975 in the Banana River, and in the Indian River near Haulover Canal and near Sebastian Inlet. At the Banana River station Sarsiella zostericola was collected in June 1974, but not again until February 1975. It was present from February to May 1975, peaking in April, but was not present in July 1975, when a single juvenile of Parasterope sp. indet. was collected (Fig. 2). At the station near Haulover Canal, Sarsiella zostericola was collected in June 1974, but the species was not again encountered until March 1975. It was present from March to July 1975, peaking in May. Sarsiella disparalis was present in small numbers from June 1974 to November 1974 (absent in December 1974), then increased in numbers starting in February, and reached a peak in May 1975. In June 1975, it had de- clined to the February number (Fig. 3). Ostracodes were extremely sparse at the station near Sebastian Inlet. One specimen of Sarsiella zostericola and 1 specimen of S. disparalis were collected in November 1975, and 1 specimen of Asteropteron oculitristis was collected in December 1975 (Fig. 4). Reproduction Ovigerous females of Sarsiella disparalis and Sarsiella zostericola were observed during the months of February, March, April, May, June, July, August (S. disparalis only), November and December. The absence of ovigerous females during other months probably reflects the few speci- mens collected during those months. Ovigerous females of other species were not collected. Adult males of myodocopid ostracodes are generally less abundant than VOLUME 90, NUMBER 4 793 S %o T °C 100 DEPTH cm e SARSIELLA ZOSTERICOLA © SARSIELLA DISPARALIS NUMBER OF SPECIMENS 1974/1975 MONTHS Fig. 2. Banana River: Seasonal changes in salinity, temperature, water depth and ostracode abundance. The number of specimens on the graph is the total num- ber collected at each station. Black dots indicate that a sample was taken during that month as well as indicating the number of specimens of Sarsiella zostericola. adult females. Two adult males of Sarsiella zostericola were collected in March 1975, from the Haulover Canal station in the Indian River. One adult male of Asteropteron oculitristis was collected in November 1974, near Sebastian Inlet, Indian River. 794 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON ate) 100 DEPTH cm fo) oO 15 © PARASTEROPE SP. @SARSIELLA ZOSTERICOLA NUMBER OF SPECIMENS I974/1975 MONTHS Fig. 3. Indian River near Haulover Canal: See legend of Fig. 2. Feeding Habit The myodocopids collected in the Indian River complex are either preda- tors or filter feeders. All known members of the genus Sarsiella are preda- tors; thus only predatory myodocopids are present in the Banana River and in the northern part of the Indian River (an exception is the single juvenile collected in the Banana River of Parasterope species indeterminate, which is a filter feeder). In the vicinity of the inlets both predatory sarsiellids and the filter feeding Asteropteron oculitristis are present. Off- VOLUME 90, NUMBER 4 795 e SARSIELLA ZOSTERICOLA a ° SARSIELLA DISPARALIS 0 ASTEROPTERON OCULITRISTIS NUMBER OF SPECIMENS (e) ow J J A Ss O N D J F M A M J J 1974/1975 MONTHS Fig. 4. Indian River near Sebastian Inlet: See legend of Fig. 2. shore, on the continental shelf, 2 filter feeders (Cycloleberis americana, Actinoseta chelisparsa) and a detritus feeder (Pseudophilomedes ferulanus) were collected. Predatory sarsiellids also are known to live offshore but were not obtained in the present study. Discussion Within the collecting area, assemblage composition and diversity appear to reflect the degree of exchange of lagoonal and oceanic waters. In the Banana River, essentially a landlocked lagoon, where exchange with oceanic water is at a minimum, only 1 species, Sarsiella zostericola, is common. 796 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON In the Indian River 2 species are common, S. zostericola and S. disparalis, but in the vicinity of inlets, where oceanic water has its greatest influence, a 3rd species, Asteropteron oculitristis, occasionally appears. The diversity of the Indian River complex is low compared to that on the continental shelf. Only 1 specimen was in each of the 3 shelf samples in the collection, but the 3 specimens represent 3 different species, thus reflecting the higher diversity of the shelf, which according to Darby (1965) contains 17 species in the vicinity of Georgia. None of the shelf species in the present collection is present in the Indian River complex, indicating that the major difference in the study area is between the Indian River complex assemblage and continental shelf assemblage. Ostracode abundance increased during the spring in both the Banana River and in Haulover Cove suggesting that a spring bloom of vegetation may be a factor, but data on vegetation are not available. Comparison of changes in salinity, temperature and water depth, with the abundance curve of the ostracodes in Figs. 2 and 3 suggests that increasing salinities during the spring may be a factor. Young, Buzas, and Young (1976) have shown that predators have a major effect on macrobenthos density in the study area; possibly, meiobenthos is also effected. Clearly, the factors determining ostracode abundance require further study. Acknowledgments I am grateful to the personnel of the Harbor Branch Foundation and the Fort Pierce Bureau of the Smithsonian Institution who collected the specimens upon which this study is based. I am especially grateful to John E. Miller, III for supplying information concerning sampling pro- cesses and station data. I thank Dr. David K. Young and Mrs. Anne Cohen for criticizing the manuscript. Literature Cited Darby, D. G. 1965. Ecology and taxonomy of Ostracoda in the vicinity of Sapelo Island, Georgia. Report no. 2 in Four Reports of Ostracod Investigations, 77 pp., 11 figures, 33 plates. Ann Arbor, Mich.: University of Michigan. [Offset report. ] Eilers, William L. 1975. Scientists report on ecological study of Indian River. Smithsonian Institution Research Reports 11:4,5, 2 text-figs. Smithsonian Institu- tion Office of Public Affairs, Washington, D.C. Young, nee K. 1976. The Indian River study. See Technology 17(10):14-18, 2 figs. . (ed.) 1975. Indian River coastal zone study annual report 1974-1975, vol. 1:180 pp., vol. 2:98 pp. Harbor Branch Consortium, Fort Pierce, Florida. . (ed.) 1976. Indian River coastal zone study annual report, 1975-1976, VOLUME 90, NUMBER 4 797 vol. 1:187 pp.; vol. Florida. Young, David K., R. M. Avent, L. I. Briel, N. J. Eiseman, R. H. Gore, R. S. Jones, G. A. Kerr, H. H. Serbert, O. von Zweck, and J. R. Wilcox. 1974. Indian River study: annual report, 1973-1974, vol. 1:183 pp., vol. 2:385 pp. Harbor Branch Consortium, Fort Pierce, Florida. Young, David K., Martin A. Buzas, and Martha W. Young. 1976. Species densities of macrobenthos associated with seagrass: A field experimental study of predation. Journal of Marine Research 34(4):577-592, 2 figs. 2:105 pp. Harbor Branch Consortium, Fort Pierce, PROC. BIOL. SOC. WASH. 90(4), 1977, pp. 798-807 BUCEROEMERSONIA, A NEW GENUS OF ISCHNOCERAN MALLOPHAGA FOUND ON THE HORNBILL GENUS TOCKUS (BUCEROTIDAE) Robert E. Elbel Abstract—Elbel, Robert E., 1518 Evergreen, Salt Lake City, Utah 84106. —Descriptions and illustrations are presented for the new genus Bucero- emersonia type clarkei and for two new species, B. clarkei and B. brelihi, found on the Hornbill genus Tockus. The distribution of three genera of Hornbill lice on Tockus is tabulated and discussed; it is suggested that B. clarkei may have evolved first from the stock that gave rise to the Hombill Ischnocera. Elbel (1976) described the new genus Bucerocophorus for three species that could not be included in either Buceronirmus or Paroncophorus. An- other new genus, Buceroemersonia, is described here for two new species, B. clarkei and B. brelihi, which were first thought to belong to a species- group of Buceronirmus but characters of the preantennal region of the head and female terminal segments precluded this possibility. Dr. K. C. Emerson, Smithsonian Institution (USNM), concurred with this opinion. For the loan of specimens appreciation is expressed to: Dr. Theresa Clay, British Museum (Natural History) (BMNH), Dr. J. A. Ledger, South African Institute of Medical Research (SAIMR), and Dr. Savo Brelih, Yugoslavian Prirodoslovni Musej Slovenije (PMS). I obtained dried mate- rial from museum skins at the USNM, American Museum of Natural His- tory (AMNH), and the Field Museum of Natural History (FMNH). Special thanks are extended to: Dr. L. T. Nielsen, Department of Biology, Univer- sity of Utah, for providing space and facilities for this study and Drs. Mary LeCroy, AMNH, C. Moreby, BMNH, M. A. Traylor, FMNH, and G. E. Watson, USNM, for supplying host and locality information. Mallophagan terminology follows Clay (1958) except as noted and nomenclature of the hosts is that of Sanft (1960). Buceroemersonia, new genus Bigs al pa Marginal carina of forehead with a short rounded dorsomedial extension posteriorly, with a long dorsal seta each side at point of nearly complete interruption laterally (dorsal submarginal seta, Clay, 1951), and with 2 long dorsal setae posterior to marginal carina (anterior dorsal setae, Clay, 1951). Ventral carina complete. Antennae filiform, similar in both sexes. Prono- tum with a long seta on each lateral margin. Pterothorax with posterior VOLUME 90, NUMBER 4 799 marginal row of long setae and a short seta on each lateral margin. Meta- sternal plate triangular, expanded posteriorly, with 6-10, normally 8, setae. Abdominal segments with tergites, sternites, and pleurites, latter not pro- longed posteriorly and with setae on margins of IV-X. Tergites H-V or II-VI, often X, in male and II-VII in female divided medially, remainder complete; on JII-VIII both sexes with a posterior marginal row of setae, post-spiracular setae being most laterad on III-VII. Sternites II-VI in male and II or II-VI in female complete and on II-VII both sexes with a posterior marginal row of setae. Normal chaetotaxy of terminal segments illustrated for each species; male X with 1-2, normally 1, ventrolateral setae each side posterior to pleural stetae and sternites VII-XI fused into single genital plate with or without indentation between VII and VIII; female sternites VII-VIII may be fused into single genital plate and spermathecal sclerite may be present; vulval margin with a row of short setae and 4-6 minute setae; postvulval sclerite long, varying in shape for each species; and fused IX—XI with 2 dorsal and 5-6 sternal setae each side. Buceroemersonia resembles most closely Buceronirmus in the dorsome- dial extension of the marginal carina which is long in Buceronirmus but short and rounded in Buceroemersonia. The dorsal submarginal setae are absent and the anterior dorsal setae are short in Buceronirmus but these setae are long in Buceroemersonia. In the row of short setae on the female vulval margin, there are minute setae in Buceroemersonia but spines in Buceronirmus. On each lateral margin of fused IX—XI in the female, sternal setae each side are at least 8 arranged vertically in Buceronirmus but 5-6 arranged laterally in Buceroemersonia. Species can most easily be separated by the shape of the ventral carina of the forehead, the number of ventral pleural setae on segments ITV—VIII, the male genitalia, and the shape of the female vulval margin. Etymology.—This genus is named for Dr. K. C. Emerson, USNM, in appreciation for his encouragement and many helpful suggestions on mal- lophagan taxonomy and on preparation of the Hornbill Mallophaga for publication. Type species.—Buceroemersonia clarkei, new species. Buceroemersonia clarkei, new species Figs. 2, 4-7 Type host.—Tockus hemprichii (Ehrenberg). Both sexes are slightly larger than corresponding sexes of B. brelihi, n. sp. only in total length. Male.—Normal chaetotaxy as in Fig. 4. Forehead with ventral carina evenly rounded anteriorly. Pterothorax with posterior marginal row of L0- 12 long setae. Abdominal tergites I-VI divided medially; remainder com- 800 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Wetaovaal. / (ASH ANY 4 fxg aus LA als Vl Bank LH re\\ 3 | leat WAT ne VOLUME 90, NUMBER 4 801 plete. Tergocentral setae: range on II, VII-VIII 6-8; III 10-12; IV 10-14; V 8-12; VI 6-10. Pleural setae each side: dorsally on IV-VII and X 1; ven- trally on V 0-1; VI-VII 1; VIII 2; X 2-4. Sternal setae: range on II 4-6; IlJ-IV 6-12; V-VI 6-10; VII 6-8; VIII 2-4. Genitalia as in Fig. 2; penis with sclerites pointed and separated anteriorly; endomeres with paired lateral lobes expanded apically; parameres angled on each posterolateral margin. Terminal segments as in Fig. 5; tergite X nearly divided medially with 4 tergocentral setae; XI with 4-6 dorsal setae and ventral margin with 6-10 setae; genital plate without indentation between VII and VIII. Female.—Sternites II-VI complete as for male. Normal chaetotaxy as in Fig. 7, differs from male as follows. Tergocentral setae: range on II 6-10; VIII 4-8. Sternal setae: range on II 4-8; IV 8-12; VII 6-10. Terminal segments as in Fig. 6; sternites VII-VIII fused into single genital plate; spermathecal sclerite long and slender, diverging posteriorly on each side of opening of spermathecal tube (Fig. 6a); vulval margin bluntly angled posteriorly with 16-22 short setae; postvulval sclerite each side long and inwardly curved. Measurements in mm.— Width Length Head Pterothorax Abdomen Head Total Male 0.36—0.39 ().34—0.38 ().46-0.57 ().46—0.50 125921576 Female 0.37-0.44 0.35-0.43 0.51-0.67 0.47-0.55 1.73-2.10 Type-material—Holotype 4, allotype °, Tockus hemprichii, USNM skin, Bodessa, Ethiopia, Africa, 20 May 1912, Childs Frick, in USNM; para- types: 1é 322, with same data; 16 422, USNM skins, Ethiopia, Africa, Feb.-May 1912, Childs Frick; 36 6 222, FMNH skin, Mt. Albana, Ethi- opia, Africa, 25 Nov. 1926, L. A. Fuertes; 16 222, FMNH skin, Alghe, Ethiopia, Africa, 23 Sep. 1941, C. W. Benson. Other specimens.—7 8 é 202 2, T. n. nasutus (Linnaeus), Uganda, Africa, Apr. 1936, Meinertzhagen 7755, BMNH; 1é 12, Cameroon, Africa, 23 Jun. 1959, J. Mouchet, BMNH; 222, FMNH skins, Sudan, Africa, 1948-1950, H. Hoogstraal; 1¢ 222, T. n. forskalii (Ehrenberg), Aden, Saudi Arabia, Nov.—Dec. 1948, Meinertzhagen, BMNH; 5¢ 6 422, T. n. epirhinus (Sun- < Figs. 1-9. Dorsal—ventral views of head and body drawn to same scale: 1, Bucero- emersonia brelihi, male head; B. clarkei—4, Entire male; 7, Entire female. Male genitalia, ventral views drawn to same scale: 2, B. clarkei; 3, B. brelihi. Terminalia, dorsal-ventral views drawn to same scale: B. clarkei—5, Male; 6, Female, a, sperma- thecal sclerite on each side of opening of spermathecal tube; B. brelili—sS, Male; 9, Fe- male. 802 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON devall), Zambia, Africa, 1953-1955, BMNH; 14 32 2, Mabelikwa, Transvaal, South Africa, 18 Jan. 1957, F. Zumpt, BMNH; 222, USNM skins, Lake Upemba, Mabwe, Zaire, Africa, Dec. 1948, G. F. de Witte; 36 6 629, T. monteiri Hartlaub, Windhoek, Namibia, Africa, 18 Jan. 1970, F. Zumpt, SAIMR; 36 6 42°, USNM skins, Waterberg, Namibia, Africa, May-Jun. 1936, W. Hoesch; 16 222, T. p. pallidirostris (Finsch and Hartlaub), Cuango River, Cafunfo, Angola, 12 May 1971, S. A. Peles, SAIMR; 1¢, USNM skins, Mpata Hills, Zambia, Africa, Apr.—Jul. 1953, E. L. Haydock; 12, T. p. neumanni (Reichenow), FNMH skin, Morogoro, Tanzania, Africa, 19 May 1917, V. G. L. van Someren; 1¢ T. pallidirostris subsp., FMNH skins, Lundazi and Kafue River, Chilula, Zambia, Africa, 1945- 1947, E. L. Button; 2¢4 12, T. a. geloensis (Neumann), Mpata Hills, Zambia, Africa, 1951, E. L. Haydock, BMNH; 6é ¢ 62 2, Muliashi, Zambia, Africa, 1952-1955, E. L. Haydock, BMNH; 44 6 22 2, AMNH skins, Ma- rungu Mts., Zaire, Africa, Mar. 1929, Rockefeller and Moses; 1é¢ 22 9, AMNH skin, T. a. suahelicus (Neumann), Monkey Bay, Malawi, Africa, 19 Sep. 1895, P. Rendall; 12, USNM skins, Nairobi, Kenya, Africa, Aug.—Nov. 1909, Loring and Mearns; 24 6 12, T. alboterminatus australis (Roberts), AMNH skins, Zululand, Natal, South Africa, Nov.-Dec. 1903, C. H. B. Grant; 1¢6 12, T. bradfieldi (Roberts), FMNH skins, Maun, Botswana, Africa, May-Jun. 1930, Vernay, Lang, and Roberts; 12, T. griseus ginga- lensis (Shaw), USNM skins, Embilipitiya, Ceylon, Jan.—Feb. 1944, S. D. Ripley; 16, USNM skin, Ceylon, B. H. Swales; 36 ¢ 622, T. birostris (Scopoli), Nepal, Mar. 1937, Meinertzhagen 9331, BMNH; 22°, FMNH skins, Madras, India, Jan. 1937, W. Koelz; 16, FMNH skin, Londa, Mysore, India, 21 Jan. 1938, W. Koelz; 3° 2, FMNH skins, Kalnahi, Uttar Pradesh, India, Feb. 1947, W. Koelz; 12, FMNH skin, RamanujGanj, Madhya Pra- desh, India, 8 Nov. 1947, W. Koelz; 46 6 12, FMNH skins, Simra, Nepal, 4 Mar. 1947, W. Koelz and R. Chand; 164 6 142 9, FMNH skins, Kotla, Kangra, Punjab, India, 1946 and 1948, W. Koelz; 56 ¢ 822, FMNH skins, Baihar, Madhya Pradesh, India, Jan.—Feb. 1949, R. L. Flemming; 48 2, FMNH skins, Hardwar, Uttar Pradesh, India, Feb. 1951, R. L. Flemming; 222, USNM skins, India, 1898, 1946-1948. Etymology.—This species is named for Dr. J. F. G. Clarke, USNM, in appreciation for financial help in collecting Oriental Mallophaga and in publishing Hornbill Amblycera (Elbel, 1967). Buceroemersonia brelihi, new species lakes; IL, 8) ce) Type host——Tockus deckeni (Cabanis). Both sexes are slightly smaller than corresponding sexes of B. clarkei only in total length. Male.—Forehead with ventral carina pointed medially (Fig. 1). Ptero- VOLUME 90, NUMBER 4 803 thorax with posterior marginal row of 12-16, normally 14, long setae. Ab- dominal tergites II-V, often VI, divided medially; remainder complete. Tergocentral setae: range on II and VIII 8-12, normally 10; III-V 12-16, normally 14; VI 10-14, normally 12; VII 10-12. Pleural setae each side: dorsally on IV-VII and X 1; ventrally on IV 0-1, normally 1; V 1-2, nor- mally 1; VI-VII 2; VII 3; X 2-3. Sternal setae: range on I, VI-VII 6-10, normally 8 on II and VII, 10 on VI; III 10-12, normally 10; IV-V 10-14, normally 12; VIII 2. Genitalia as in Fig. 3, penis rounded and closed anteriorly; endomeres with paired lateral lobes narrow; parameres with a rounded knob on each posterolateral margin. Normal chaetotaxy of ter- minal segments as in Fig. 8; tergite X divided medially with 2 tergocentral setae; XI with 6-8 dorsal setae and ventral margin with 4-6 setae. Female.—Sternite II complete, III-VII circular or oval in shape. Chaeto- taxy differs from male as follows. Tergocentral setae: range on II and VIII 10-12. Sternal setae: range on III 10-14, normally 12; VI 10-12. Normal chaetotaxy of terminal segments as in Fig. 9; sternite VII separate from VIII; spermathecal sclerite at most minute extensions on each side of spermathecal tube; vulval margin evenly rounded posteriorly with 12-18 short setae; postvulval sclerite each side long and outwardly curved with anterolateral projection. Measurements in mm.— Width Length Head Pterothorax Abdomen Head Total Male 0.36-0.41 Ors22030 0).47-0.58 ().42-0.48 oes Female 0.41-0.44 0.37—0.40 0.57-0.65 0.48—-0.52 1.63-1.80 Type-material—Holotype ¢, allotype ?, Tockus deckeni, USNM skins, Tana River, Kenya, Africa, Aug. 1912, Childs Frick, in USNM; paratypes: 766 1222, with same data; 76 ¢ 722, USNM skins, Kenya, Africa, Jul.— Aug. 1912, Childs Frick; 1¢ 12, Koka, Ethiopia, Africa, 13 Dec. 1960, S. Brelih, PMS; 16 12, USNM skins, Gato River, Gardula, Ethiopia, Africa, Apr.—May 1912, Childs Frick; 5¢ 6 229, USNM skins, Ethiopia, Africa, Jun. 1912, Childs Frick. Other specimens—1é 422, T. nasutus forskalii (Ehrenberg), Aden, Saudi Arabia, Nov. 1948, Meinertzhagen 17753, BMNH; 10¢ ¢ 722, T. e. erythrorhynchus (Temminck), Somalia, Africa, Jan—Feb. 1949, Meinert- zhagen, BMNH; 24 ¢ 22 2, Marsabit, Kenya, Africa, 21 Jan. 1956, Meinert- Zhagen 20512, BMNH; 7¢64 322, FMNH skins, Ethiopia, Africa, Dec. 1926, A. M. Bailey; 26 6 322, USNM skins, Ethiopia and Kenya, Africa, Jun.—Jul. 1912, Childs Frick; 12, T. e. rufirostris (Sundevall), Gravelotte, Transvaal, South Africa, 28 Oct. 1957, F. Zumpt, BMNH; 3¢ ¢ 22 2, Goha Hill, Botswana, Africa, 21 Sep. 1958, F. Zumpt, BMNH; 1é 1°, Chirundu, 804 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Zambia, Africa, 17 Feb. 1964, F. Zumpt, BMNH; 32 2, Kariba, Rhodesia, Africa, 16 Feb. 1964, F. Zumpt, SAIMR; 346 6 82 2, T. f. flavirostris (Rup- pell), Hargeisa, Somalia, Africa, Jan—Feb. 1949, Meinertzhagen, BMNH; 266 12, Isiolo, Kenya, Africa, Feb. 1956, Meinertzhagen 20455, BMNH; 2648, USNM skins, Ethiopia, Feb.Jun. 1912, Childs Frick; 16, T. f. leuco- melas (Lichtenstein), Limpopo River, Chicualacuala, Mozambique, Africa, 1] Jul. 1953, H. E. Paterson, BMNH; 36 ¢ 32 2, Debeeti, Botswana, 18 Jul. 1956, F. Zumpt, BMNH; 1é, Newington, Transvaal, South Africa, 17 Jul. 1957, F. Zumpt, BMNH; 14, T. alboterminatus geloensis Xx suahelicus, AMNH skin, Meru Forest, Kenya, Africa, 10 Aug. 1912, Childs Frick; 2 ¢, USNM skins, with same data. Etymology.—This species is named for Dr. Savo Brelih, PMS, in appre- ciation for the loan of this type-material as well as other Hornbill Ambly- cera and Ischnocera (Elbel, 1967 and 1976). | Discussion Both species are similar in size, Buceroemersonia clarkei being slightly larger than corresponding sexes of B. brelihi only in total length. The ven- tral carina of the forehead is evenly rounded anteriorly in B. clarkei but pointed medially in B. brelihi (Figs. 1 and 4). The posterior marginal row on pterothorax has at most 12 long setae in B. clarkei but at least 12 in B. brelihi. There are normally more tergocentral setae on abdominal seg- ments II-VIII of B. brelihi than corresponding sexes of B. clarkei but on X of males, B. brelihi has 2 tergocentral setae and B. clarkei has 4 (Figs. 5 and 8). Both species have a dorsal pleural seta on each side of IV—VII and X (Figs. 4, 7, 8-9). Ventrally on each side of IV-VIII B. brelihi nor- mally has a pleural seta on IV-V, 2 on VI-VII, and 3 on VIII (Figs. 8-9) but B. clarkei has 1 on VI-VII and 2 on VIII (Figs. 5-6). Sternal setae on II and IV-VI are normally more numerous in B. brelihi than in correspond- ing sexes of B. clarkei. In the male genitalia penial sclerites are pointed and separated anteriorly only in B. clarkei which is also unique in having endomeres with the paired lateral lobes expanded apically and parameres that are angled on each posterolateral margin (Fig. 2). In B. brelihi the penis is rounded and closed anteriorly, the paired lateral lobes of endomeres are narrow, and the parameres have a rounded knob on each posterolateral margin (Fig. 3). The male tergite X is divided medially and the fused male genital plate is indented between VII and VIII only in B. brelihi (Fig. 8). There are more setae on tergite XI in males of B. brelihi than B. clarkei but the reverse is true for setae on the ventral margin (Figs. 5 and 8). Female sternites III-VII are circular or oval in shape for B. brelihi (Fig. 9) but II-VI are complete in B. clarkei (Fig. 7) and sternites VII-VIII are fused into a single genital plate only in B. clarkei (Figs. 6 and 9). Female sperma- thecal sclerite with long slender margins diverging posteriorly on each side of opening of spermathecal tube is present only in B. clarkei (Fig. 6a) > VOLUME 90, NUMBER 4 805 Table 1. Distribution of Buceroemersonia, Bucerocophorus, and Chap- inia (Elbel, 1967, 1976) on the genus Tockus (Bucerotidae) arranged ac- cording to Sanft (1960). Host Buceroemersonia Bucerocophorus Chapinia Tockus n. nasutus B. clarkei C. lophocerus T. n. forskalii B. clarkei B. brelihi T. n. epirhinus B. clarkei C. lophocerus T. n. dorsalis T. e. erythrorhynchus B. brelihi C. lophocerus T. e. rufirostris B. brelihi C. lophocerus T. e. damarensis T. deckeni B. brelihi C. lophocerus T. f. flavirostris B. brelihi C. lophocerus T. f. somaliensis T. f. elegans T. f. leucomelas B. brelihi C. lophocerus T. hemprichii B. clarkei T. monteiri B. clarkei T. p. pallidirostris B. clarkei B. latifrons T. p. neumanni B. clarkei T. c. camurus C. camuri T. c. pulchrirostris T. h. hartlaubi T. h. granti C. fasciati T. f. fasciatus B. latifrons C. fasciati T. f. semifasciatus B. latifrons T. a. alboterminatus B. latifrons T. a. geloensis B. clarkei B. latifrons C. fasciati T. a. geloensis * suahelicus B. brelihi T. a. suahelicus B. clarkei B. latifrons C. fasciati T. alboterminatus australis B. clarkei B. latifrons C. fasciati T. bradfieldi B. clarkei T. g. griseus C. clayae T. griseus gingalensis . Clarkei T. birostris . Clarkei C. clayae jeoles) posteriorly the female vulval margin is bluntly angled in B. clarkei but evenly rounded in B. brelihi and there are normally more setae on this mar- gin in B. clarkei than in B. brelihi; the long postvulval sclerite each side is inwardly curved in B. clarkei but outwardly curved with an anterolateral projection in B. brelihi (Figs. 6 and 9). Aviparasitological Relations Species of the ischnoceran genus Buceroemersonia infest Hornbill hosts only in the genus Tockus and all species are infested except 7. camurus, 806 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON T. hartlaubi, and T. fasciatus (Table 1). B. clarkei was found alone on T. hemprichii, T. monteiri, and T. bradfieldi, in association with Buceroco- phorus latifrons on T. pallidirostris, and with Chapinia clayae on T. griseus and T. birostris. The only ischnoceran species found on T. erythrorhynchus, T. deckeni, and T. flavirostris was Buceroemersonia brelihi but a few B. brelihi were found on T. nasutus and T. alboterminatus, host species in- fested by B. clarkei. In fact on T. n. forskalii both B. clarkei and B. brelihi infested the same host specimen. C. lophocerus shared common hosts with these ischnoceran lice except on T. alboterminatus where C. fasciati and Bucerocophorus latifrons were found. Tockus is the only Hornbill genus with members in both the Ethiopian and Oriental Regions (Sanft, 1960). As noted by Elbel (1967), the ambly- ceran, C. clayae, found on the Oriental species, T. griseus and T. birostris, resembled most closely C. acutovulvata, found on Oriental species of Anthracoceros, rather than Chapinia that infested Ethiopian species of Tockus. Kellogg (1896) stated that Mallophaga spent their entire lives on the host bird and new hosts were infested by migration of lice from one bird to another during copulation, nesting, or roosting. According to Clay (1949), birds of different species normally did not come in close enough contact to transfer lice. However, Clay (1962) described natural straggling as occurring between hosts that happened to be nesting in close proximity and establishment on the new host might be facilitated by the absence of a resident louse. Elbel (1967) suggested that there may have been more recent contact between the Indian Tockus and Anthracoceros whose ranges overlapped than between the more nearly related Indian and African Tockus; natural straggling may have accounted for establishment of Chapinia on Anthracoceros. This view is not supported by the ischnoceran Buceroemersonia clarkei which was found on Tockus in both regions, but speciation rates in the Amblycera and Ischnocera have been so different that comparisons on the same host group have little value (Clay, 1957). According to Kellogg (1896) the ancestral bird species spread and gave rise to geographical races which eventually became distinct species, often dis- tinguished only by superficial differences in color, etc., but the Mallophaga remaind nearly the same because their environment was essentially the same. The environment of the Mallophaga, the physical and chemical composition of the feathers and blood, presumably changes slower than do other factors leading toward speciation of the bird (Clay, 1949). For ex- ample, B. clarkei infests 13 hosts in 8 species in both the Ethiopian and Oriental regions but B. brelihi infests 7 hosts in 5 species in the Ethiopian region. Female sternites II-VI are complete in B. clarkei (Fig. 7) as for males of Buceroemersonia, Bucerocophorus and Buceronirmus, but III-VII are circular or oval in shape for B. brelihi (Fig. 9) as for females of Bucero- cophorus (Elbel, 1976) and Buceronirmus (Elbel, 1977). With its wider VOLUME 90, NUMBER 4 807 distribution and uniqueness of the female abdominal sternites, B. clarkei may have evolved first from the stock that gave rise to the Hornbill Ischno- cera. Similarly, Elbel (1967) thought that both Ethiopian and Oriental species of Chapinia shared a common ancestor of Chapinia on Tockus be- fore the Indian and African Tockus became separated, but once separated the Chapinia evolved as did the Hornbills to the recognized species within each region. Literature Cited Chapin, J. P. 1954. Gazetteer for “The birds of the Belgian Congo.” Bull. Amer. Mus. Nat. Hist. 75B:638—738. Clay, T. 1949. Some problems in the evolution of a group of ectoparasites. Evolu- tion 3:279-299. 1951. An introduction to a classification of the avian Ischnocera (Mallo- phaga): Part I. Trans. R. Ent. Soc. London 102:171—194. 1957. The Mallophaga of birds. Pages 120-158 in First symposium on host specificity among parasites of vertebrates. JI.U.B.S. and Univ. Neuchatel, Neuchatel. 1958. Revisions of Mallophaga genera. Degeeriella from the Falconiformes. Bull. British Mus. (Nat. Hist.) Ent. 7:121—207. 1962. A key to the species of Actornithophilus Ferris with notes and de- scriptions of new species. Bull. British Mus. (Nat. Hist.) Ent. 11:191—252. Elbel, R. E. 1967. Amblyceran Mallophaga (biting lice) found on the Bucerotidae (Hornbills). Proc. U.S. Natl. Mus. 120:1-76. ——. 1976. Bucerocophorus, a new genus of ischnoceran Mallophaga from African Hombills (Bucerotidae). Proc. Biol. Soc. Washington 89:313-324. ——. 1977. Two new Buceronirmus (Mallophaga: Philopteridae) from Rhyticeros undulatus and R. plicatus (Hornbills). Pacific Ins. 17:413-418. Friedmann, H. 1930. Birds collected by the Childs Frick expedition to Ethiopia and Kenya colony. Part I. Non-Passeres. U.S. Natl. Mus. Bull. 153:xiii + 516 pp. Kellogg, V. L. 1896. New Mallophaga I: With special reference to a collection made from maritime birds of the bay of Monterey, California. Proc. California Acad. Sci. 6:31-182. Sanft, K. 1960. Bucerotidae (Aves: Upupae). Das Tierreich 76:1—176. PROC. BIOL. SOC. WASH. 90(4), 1977, pp. 808-812 THE TYPES AND TYPE-LOCALITIES OF OREORTYX PICTUS (DOUGLAS) AND ORTYX PLUMIFERUS GOULD M. Ralph Browning National Fish and Wildlife Laboratory, U.S. Fish and Wildlife Service, National Museum of Natural History, Washington, D.C. 20560. There are two northern races of the North American species known as Oreortyx pictus (Mountain Quail). According to the fifth edition of the A.O.U. Check-list of North American birds (1957), the dark form on the west side of the Cascade Mountains from northwestern California north- ward to western Washington is O. pictus palmeri Oberholser, 1923. The other, a pale race, is found east of the Cascade Mountains in eastern Wash- ington southward to east-central California, and is called O. p. pictus (Douglas, 1829). Oreortyx plumiferus (Gould, 1837) was synonymized with Oreortyx pictus pictus by Oberholser (1923). A critical appraisal of the pertinent literature and the existing types re- veals that the name plumiferus should be applied to the pale form and pictus to the dark form. This is a return to the treatment in the third (1910) and earlier editions of the A.O.U. Check-list. The remainder of this paper documents the evidence supporting this arrangement. The specimens upon which the names pictus and plumiferus were based were collected by David Douglas during his travels in western North America. Originally regarded as species of Ortyx, these names were first written as picta and plumifera. However, Ortyx (and Oreortyx) is not femi- nine but masculine, according to Woods (1944: 170), and the termination of the trivial names has been changed accordingly. In the fall of 1826, Douglas traveled from Ft. Vancouver (=Vancouver, Washington), southward along the Multnomah (=Willamette) River to near what is now Eugene in Lane County, Oregon, thence southwestward to the junction of Elk Creek and the Umpqua River, and further south to near what is now Roseburg (Douglas, 1914; Morwood, 1973). Careful scrutiny of Douglas’s activities, particularly as reported in his journal (Douglas 1914:220) has revealed that he collected a specimen of O. pictus northeast of what is now Elkton in western Douglas County on 12 October. He wrote that the specimen was too badly shattered by shot to be pre- served. Although Douglas made no entries in his journal that would sug- gest he collected additional specimens of pictus during October or Novem- ber, he later stated (Douglas, 1829b:145) that he preserved specimens in November 1826 but failed to mention where these specimens were col- VOLUME 90, NUMBER 4 309 lected. Nevertheless, according to his journal (Douglas, 1914), he must have been somewhere between Elkton and Eugene or further north along the Willamette River. Douglas (1829b) reported losing these specimens while crossing a river in November 1826. Such a mishap occurred on 15 November (Douglas, 1914:237) while crossing the Sandiam (=Santiam) River, a tributary of the Willamette River in Linn County, Oregon. Douglas also traveled in California (Morwood, 1973); but his activities there are not well known because his journals and many specimens were lost in another accident on the Fraser River, British Columbia (Jepson, 1933). Although Douglas may have collected specimens of Oreortyx pictus while in California, he did not visit there until late 1830, well after Oreortyx pictus was described (Douglas, 1829a). The A.O.U. Check-list (1931; 1957) gave the type-locality of pictus as the “interior of California as far north as 45°N”°—the range given by Douglas (1829b). As we have just seen, this is misleading since “California” in Douglas’ day also included Oregon. Oberholser (1923) concluded that the lost specimens of pictus collected by Douglas had been taken in western Oregon. Oberholser, apparently un- aware of the posthumous publication of Douglas’s journals (1904; 1914), restricted the type-locality of pictus to the headwaters of the Umpqua River near the Calapooya Mountains. Although birds from that region are of the pale form, Douglas neither visited the headwaters of the Umpqua River nor was he near the Calapooya Mountains. The only evidence concerning the provenance of the specimens on which Douglas based the name pictus comes from his statements that he shot and discarded a specimen in October and that he preserved but lost specimens in November. The October specimen is not mentioned in the original description (Douglas, 1829a) or in Douglas’s (1829b) more detailed discussion on pictus, but the November specimens are discussed in the latter publication. Although Douglas (1829a, 1829b) did not so state, the original description would appear to have been based on the November specimens. During November, Douglas was west of the Cascade Mountains where he crossed the Santiam River on 15 November 1826. Therefore, I restrict the type-locality of Oreortyx pictus to the junction of the Willa- mette and Santiam rivers, Linn County, Oregon. The name pictus is thus applicable to the dark form. Because the name palmeri Oberholser, 1923 (type-locality: Yaquina, Lincoln County, Oregon), is also applicable to this race, it becomes a subjective junior synonym. Subsequent to his description of O. pictus, Douglas obtained additional specimens. Three were discovered among Douglas's effects (Gould, 1850) and were the basis for Gould’s (1837a) “Ortyx plumifera.”. Gould (1837a, 1837b) gave the range of plumiferus as “California,” a name which at that date applied to the region south of the Columbia River. Presumably, be- cause Douglas had shipped his biological materials to London from the 810 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Hawaiian Islands in early September 1832 and the specimens available to Gould had been among Douglas’s effects, these specimens must have been acquired by Douglas between October 1832 and October 1833. During that time Douglas was in the Pacific Northwest, and from there he made his last trip to the Hawaiian Islands where he remained until his death in 1834 (McKelvey, 1955). Oberholser (1923:83) restricted the type-locality of plumiferus to the northern Willamette Valley in western Oregon “since Douglas was known to have traveled in this general region.” Oberholser (1923) believed that the plate of plumiferus in Gould (1837a) represented a bird of the pale race. Oberholser did not, however, examine specimens from the Willamette Valley (according to Allan Duvall, unpub- lished notes, U.S. Fish and Wildlife Service) until after his 1923 publica- tion. Then he decided that birds from the northern Willamette Valley be- longed to the dark form “palmeri,” to which indeed they do. Oberholser (fide Duvall’s notes) maintained that he erred only in his restriction of the type-locality of plumiferus and that Gould’s plate clearly represented the paler birds. I have examined Gould’s (1837a) plate, in addition to a later plate also by Gould (1850) and find that the dorsal coloration of the birds in both plates resembles specimens representing the pale form. The type-series of plumiferus is in the British Museum (Natural History). Specimen BM 1855.12.19.30, a dismounted bird and considered the type of plumiferus (according to Swarth, in Grinnell, 1932), probably was se- lected by Ogilivie-Grant. However, because Gould did not designate a holotype, BM 1855.12.19.30 is a lectotype. Gould (1850) stated that only three specimens were found among Doug- las’s belongings. But Derek Goodwin (in litt., 7 November 1975) of the British Museum, who was kind enough to examine the lectotype and other specimens of the presumed type-series of plumiferus, found a fourth (un- catalogued) specimen. The lectotype was catalogued in 1855; the other two specimens, in 1881. The lectotype and another specimen of the series (BM 1881.5.1.5509) resemble individuals of the pale form, whereas BM 1881.5.1. 0910 and the uncatalogued specimen resemble individuals of the dark form (Goodwin, in litt.). According to van Rossem (1937), specimens of O. pictus are vulnerable to rather rapid postmortem changes in color. However, specimens of the dark race in the National Museum of Natural History and the Academy of Natural Sciences, Philadelphia, collected in the mid-1800’s are similar in color to recently collected specimens from the same region. Although the lectotype of plumiferus was at one time mounted, there are no indications of bleaching from exposure, such as might have occurred had the specimen been on display, and which would explain the pallor of the lectotype (Goodwin, in litt.). The fact that the lectotype is a young adult (Swarth, in VOLUME 90, NUMBER 4 811 Grinnell, 1932) has no bearing since dorsal coloration of O. pictus does not vary with maturity (van Rossem, 1937). Goodwin (in litt.) found the lectotype of plumiferus to be only slightly darker than specimens of O. p. russelli Miller, 1946, from southern Cali- fornia, a race that is even paler than the pale northern form. Thus, the color of the upper parts of the lectotype of plumiferus and the plates of the species (Gould 1837a, 1850) indicate that the name plumiferus should apply to the pale form. The type-series of plumiferus may have been acquired from a hunter at Ft. Vancouver whom Douglas (1904:261, 262) had asked to collect speci- mens of O. pictus. If so, he apparently obtained birds from both slopes of the Cascade Mountains. The two dark specimens in the type-series of plumiferus must have been collected within the range of nominate pictus (the dark race) whereas the lectotype of plumiferus and BM 1881.5.1.5509 were no doubt collected from the east slope of the Cascades. Travel from Ft. Vancouver (west of the Cascades) eastward to the interior was not un- common in Douglas's time. From Ft. Vancouver the nearest population of the pale form is in Wasco County, Oregon, approximately 90 air miles dis- tance to the east and to which I hereby restrict the type-locality of plumiferus. In summary, the dark northern race of Mountain Quail should be known as Oreortyx pictus pictus (restricted type-locality: Junction of the Willamette and Santiam rivers, Linn County, Oregon), which occurs west of the Cas- cade Mountains. The pale northern race found east of the Cascades should be known as O. p. plumiferus (restricted type-locality: Wasco County, Oregon). The name O. p. palmeri is a subjective synonym of O. p. pictus. This treatment, insofar as pertaining to the nomenclature and ranges of the two races, follows Ridgway (in Baird et al., 1874) and the first three editions of the A.O.U. Check-list. Therefore, palmeri of the A.O.U. Check- list (1957) becomes a synonym of pictus, and pictus (sensu A.O.U., 1957 reverts to plumiferus. Acknowledgments For the opportunity to examine specimens I am indebted to the curators of the following collections: American Museum of Natural History, Car- negie Museum of Natural History, Museum of Comparative Zoology, Oregon State University Museum of Natural History, Philadelphia Acad- emy of Natural Sciences, Portland State University, and University of Puget Sound. Access to Gould’s plates was provided by the Philadelphia Academy of Natural Sciences. I am especially grateful to Derek Goodwin for comparing critical specimens in the British Museum (Natural History) and for providing information on the type-series of plumiferus. 1 thank 812 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON J. W. Aldrich, E. Eisenmann, A. L. Gardner, D. Goodwin, and S. Olson for their criticisms of earlier drafts of this paper. Literature Cited American Ornithologists’ Union. 1910. Check-list of North American birds. Third edition. Amer. Omithol. Union, New York. ———., 1931. Check-list of North American birds. Fourth edition. Amer. Omithol. Union, Lancaster. —. 1957. Check-list of North American birds. Fifth edition. Amer. Omithol. Union, Baltimore. Baird, S. F., T. M. Brewer, and R. Ridgway. 1874. A history of North American birds. Vol. 3. Little, Brown, and Co., Boston. Douglas, D. 1829a. Observations on some species of the genera Tetrao and Ortyx, natives of North America; with descriptions of four new species of the former, and two of the latter genus. Phil. Magazine, new. ser. 5:73-74. (January 1829) — —. 1829b. Observations on some species of the genera Tetrao and Ortyx, natives of North America; with descriptions of four new species of the former, and two of the latter genus. Trans. Linn. Soc. London 16:133-149. (before May 1829) 1904. Sketch of a journey to the northwestern parts of the continent of North America during the years 1824—25-26-27. Quart. Oregon Hist. Soc. 5(3):230-271. —. 1914. Joumal kept by David Douglas during his travels in North America, 1823-27. William Wesley and Sons, London. Gould, J. 1837a. Icones Avium. Privately published, London. (August 1837) 1837b. [Description of Ortyx plumifera, read 11 April 1837.] Proc. Zool. Soc. London, p. 42. (5 December 1837) 1850. A monograph of the Odontophorinae or partridges of America. Pri- vately published, London. Grinnell, J. 1932. Type localities of birds described from California. Univ. California Publ. Zool. 38:243-324. Jepson, W. L. 1933. David Douglas in California. Madrono 2:97—100. MckKlevey, S. D. 1955. Botanical Exploration of the Trans-Mississippi West 1790- 1850. Armold Arboretum Harvard Univ., Jamaica Plains, Massachusetts. Morwood, W. 1973. Traveler in a vanished landscape: The life and times of David Douglas. Clarkson N. Potter, Inc. New York. Oberholser, H. C. 1923. Notes on the forms of the genus Oreortyx Baird. Auk 40: 80-84. van Rossem, A. J. 1937. A review of the races of the Mountain Quail. Condor 39: 20-24. Woods, R. S. 1944. Naturalist’s Lexicon. Abbey Garden Press, Pasadena, California. PROC. BIOL: SOC. WASH. 90(4), 1977, pp. 813-818 THE STATUS AND SYSTEMATIC POSITION OF THE SPECIES OF THE BOPYRID ISOPOD GENUS PHYLLODURUS STIMPSON, 1857 John C. Markham Abstract.—Markham, J. C., Bermuda Biological Station for Research, Inc., St. George's West, 1-15, Bermuda.—The monotypic bopyrid isopod genus Phyllodurus Stimpson, 1857, is assigned, as type-genus and _ sole representative, to a new subfamily, Phyllodurinae, intermediate in charac- ters and mode of host selection between the subfamilies Athelginae Cod- reanu and Codreanu and Ioninae H. Milne Edwards. An annotated synonymy for the single species of Phyllodurus, P. abdominalis Stimpson, 1857, is presented. The only other nominal species of Phyllodurus, P. ro- bustus Pearse, 1953, is placed in synonymy with Pseudione upogebiae Hay, IS, The systematic position of the species Phyllodurus abdominalis Stimpson within the family Bopyridae has long been uncertain because it does not fit into any of the currently recognized subfamilies. Moreover, another species, P. robustus Pearse, was clearly incorrectly assigned to this genus, but its true identity could not be determined from the published descrip- tion. To resolve these problems, I am proposing a new subfamily for Phyllodurus abdominalis and presenting evidence that P. robustus is a junior synonym of Pseudione upogebiae Hay, 1917. Thanks are extended to Dr. Thomas E. Bowman of the National Museum of Natural History, Smithsonian Institution, for the loan of material (desig- nated USNM) and for valuable comments on the manuscript. This report Was prepared under National Science Foundation Grant number DEB 76-20102 administered through the Bermuda Biological Station for Re- search, Inc., of which this is scientific contribution number 691. Family BOPYRIDAE Ratinesque, 1815 Subfamily PHYLLODURINAE, new subfamily Diagnosis Female: All segments distinct; body axis nearly straight; frontal lamina, coxal plates and dorsolateral pereonal bosses all moderately developed; no middorsal projections on pereomeres; pleon triangular, markedly narrower than pereon, extending far posteriorly; pleon of 6 pleo- meres, first with dorsal papillose process at each side of anterior margin; pleon bearing pedunculate falcate lateral plates and pleopodal exopodites, both with entire margins, on each of first 5 pleomeres, all of nearly same 814 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON size; no pleopodal endopodites; terminal pleomere produced into sharp point, bearing uniramous uropods of same structure as lateral plates and exopodites. Male: Body more than twice as long as wide; all body regions and segments distinct; pleon of 6 pleomeres, last 5 produced into con- stricted lateral lobes extending posterolaterally; 5 pairs of uniramous sac- like pleopods; no uropods; terminal pleomere pointed posteriorly. Ab- dominal parasite of Upogebia. Type-genus, and sole representative, Phyllodurus Stimpson, 1857 Discussion —Phyllodurus Stimpson has always been difficult to place within the family Bopyridae because it has not fit into any of the estab- lished subfamilies. Shiino (1965) placed it in the “Athelges-group” (=Athel- ginae Codreanu and Codreanu), and in an unpublished treatment of the family Bopyridae, I tentatively assigned it to the subfamily Ioninae H. Milne Edwards. Clearly, its affinities are closest to these two subfamilies. It differs from all other athelgines in that their females are always notice- ably distorted, their pereomeres are often only obscurely separated, their heads are reduced, deeply embedded in their pereons and often partly fused with some pereomeres; the males of the Athelginae all completely lack pleonal segmentation and appendages; and their hosts, while infested abdominally, are always paguroids. Phyllodurus differs from all ionines in that the females of the latter are at least moderately distorted; their pleons are deeply embedded in their pereons; the pleonal appendages are all elongate and with tuberculate or digitate margins; male ionines are rela- tively more elongate, and their pleomeres are not laterally produced; and the hosts of ionines, although occasionally callianassids and upogebiids (much more often brachyurans), are infested only branchially. On the basis of morphology and host selection, Phyllodurus (and thus its subfamily Phyllodurinae) seems to represent an evolutionary link among several bopyrid subfamilies, as Shiino (1952, 1965) has pointed out. The subfamily Pseudioninae, taken as the most generalized, is largely restricted to anomuran hosts, some species of the rather primitive genus Pseudione infesting the genera Callianassa and Upogebia. The subfamily Ioninae, which shares many morphological characters with Phyllodurinae, contains mostly parasites of brachyurans, but several of the most primitive ionines, including all species of Ione, the single species of Procepon and one species of Hypocepon, infest callianassids. All members of the Pseudioninae and the Ioninae are branchial parasites, in contrast with Phyllodurus. The sub- family Phyllodurinae shares many morphological characters and the loca- tion of attachment with the Athelginae, but members of these two sub- families infest hosts in different superfamilies within the infraorder Anomura. VOLUME 90, NUMBER 4 815 Phyllodurus Stimpson, 1857 Phyllodurus Stimpson, 1857:511 [Type-species, by monotypy, Phyllodurus abdominalis Stimpson]. Diagnosis.—As for subfamily Phyllodurinae. Only one species, Phyllodurus abdominalis Stimpson. Phyllodurus abdominalis Stimpson, 1857 Phyllodurus abdominalis Stimpson, 1857:511-513 [Type-localities, Puget Sound, Washington, and Tamales Bay, California; infesting “the common Gebia’ (evidently = Upogebia pugettensis (Dana) ].—Lockington, 1877: 57 [Tomales Bay; infesting U. pugettensis; first description of male]; 1878:299-300 [Tomales Bay; infesting U. pugettensis|.—Stebbing, 1893: 418.—Calman, 1898:261, 282 [Puget Sound; host unspecified ].—Richard- son, 1899a:868; 1899b:337; 1900:309; 1904a:78 [San Francisco Bay, California; infesting U. pugettensis]; 1905:540-544; figs. 582-585.—Bon- nier, 1900:171, 215-217, 351.—Holmes, 1900:158.—Gerstacker and Ort- mann, 1901:184, 235-236, 266.—Nierstrasz and Brender a Brandis, 1923: 80; 1931:209 [Nanaimo, British Columbia; no host recorded].—Fee, 1926:39 [Departure Bay, British Columbia; infesting U. pugettensis].— Fraser, 1932:64 [Nanaimo, British Columbia, and San Juan Islands, Washington; infesting U. pugettensis]|—MacGinitie, 1935:658, 659-660, 704, 708 [Monterey Bay, California; infesting U. pugettensis|.—Hatch, 1947:164, 224; pl. IX, figs. 108, 109.—MacGinitie and MacGinitie, 1949: 265, 266, 292; fig. 120c [Tomales Bay; infesting U. pugettensis]; 1968: 265, 266, 292; fig. 120c.—Pearse, 1953:237.—Menzies and Miller, 1954: 141, 153.—George and Stromberg, 1968:251, 253 [San Juan Island, Wash- ington; infesting U. pugettensis]—Sadoglu, 1969:197.—Schultz, 1969: 321; fig. 512.—Stromberg, 1971:2 [San Juan Islands; infesting U. puget- tensis |.—Kozloff, 1974:148.—Miller, 1975: 285, 286, 305; pl. 64, fig. 16. Phyllodurus |sp.].—MacGinitie, 1937:1035. “Isopods.”—Kozloff, 1973:233. Remarks.—Phyllodurus abdominalis is known from only a moderate dis- tance along the Pacific coast of North America, from southern British Columbia to central California, and it evidently infests only a single host species, Upogebia pugettensis. Nonetheless, it is very common in parts of its range, particularly the San Juan Islands and Tomales Bay, and it has been collected many times. It has been well described and illustrated, es- pecially by Richardson (1905), so nothing need be added to its description here. 816 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Status of Phyllodurus robustus Pearse, 1953 Pearse (1953) described a parasite of Upogebia affinis (Say) from Alli- gator Harbor, Florida (Gulf of Mexico) under the name of Phyllodurus robustus. In order to determine its status, I have examined the type-speci- mens of P. robustus, USNM 93719, of Pseudione upogebiae Hay, USNM 48369 and USNM 48370 (infesting Upogebia affinis at Beaufort, North Carolina) and of Pseudione furcata Richardson, USNM 29093 (infesting an unknown host off the coast of Virginia). The original description of Phyllodurus robustus by Pearse (1953) was not adequate for one to ascertain the true nature of that species except that its assignment to the genus Phyllodurus was clearly erroneous. The type- specimens were mounted on a microscope slide and crushed and dissolved almost beyond recognition. Careful examination of them reveals most of the characters diagnostic for Pseudione upogebiae, other characters being indiscernible. Specifically, the female’s pereopods bear prominent basal carinae, and its pleopods are deeply dentate, while the male has a distinc- tive outline, and its head is very reduced and fused with the first pereo- mere. The reasons for Pearse’s assignment of this species to the genus Phyllo- durus are unclear. Phyllodurus abdominalis, as implied by its name, is an abdominal parasite, though Pearse (1953) clearly mentions that P. robustus infested its host branchially. To be sure, the host of P. abdominalis is Upo- gebia pugettensis, a congener of the host of P. robustus, U. affinis. This, however, is not in itself an adequate explanation, because Pearse (1945, 1947) twice recorded collecting Pseudione upogebiae infesting Upogebia affinis before he described Phyllodurus robustus. Hay (1917) considered Pseudione upogebiae to be closely similar to P. furcata. Having examined the types and other specimens of P. furcata, I am satisfied that there is no question about the distinctness of P. upo- gebiae from P. furcata, even though they both infest the same host species, Upogebia affinis, at the same localities. The male of P. furcata remains undescribed, but specimens of it which I have seen are also clearly distin- guishable from that of P. upogebiae. Lemos de Castro (1965) speculated that Phyllodurus robustus might be a synonym of Pseudione upogebiae. He was of course unable to confirm that without examining the types of Phyllodurus robustus because of the inadequacy of the published description. His report of Pseudione upo- gebiae from Brazil makes this one of the most widespread of all western Atlantic bopyrids. Synonymizing Phyllodurus robustus with Pseudione upogebiae makes the genus Phyllodurus once again monotypic, as is the new subfamily con- taining it, Phyllodurinae. VOLUME 90, NUMBER 4 817 Literature Cited Bonnier, J. 1900. Contribution a l’étude des épicarides. Les Bopyridae. Trav. Sta. Zool. Wimereux 8:1—476. Calman, W. T. 1898. On a collection of Crustacea from Puget Sound (Puget Sound Zoology, Columbia University Contributions no. 9). Ann. New York Acad. Sci. 11:259-292. Fee, A. R. 1926. The Isopoda of Departure Bay and vicinity, with descriptions of new species, variations and colour notes. Contrib. Canadian Biol., Toronto, new ser. 3(2):13—46. Fraser, C. M. 1932. A comparison of the marine fauna of the Nanaimo region with that of the San Juan Archipelago. Trans. Roy. Soc. Canada ser. 3, 26 (Section V—Biol. Sci.):49-70. George, R. Y., and J.-O. Stromberg. 1968. Some new species and new records of marine isopods from San Juan Archipelago, Washington, USA. Crustaceana 14: 225-254. Gerstaecker, A., and A. E. Ortmann. 1901. Crustacea, Malacostraca. Bronns Klassen und Ordnungen des Thier-Reichs 5, Abteil 1, Halfte 2, Lieferungen 1—24:i-vii, 1-688. Hatch, M. H. 1947. The Chelifera and Isopoda of Washington and adjacent regions. Univ. Washington Publ. in Biol. 10:155-274. Hay, W. P. 1917. A new genus and three new species of parasitic isopod crustaceans. Proc. U.S. Nat. Mus. 51:569-574. Holmes, S. J. 1900. Synopsis of the California stalk-eyed Crustaceae. Occas. Pap. California Acad. Sci. 7: 1-262. Kozloff, E. N. 1973. Seashore life of Puget Sound, the Strait of Georgia, and the San Juan Archipelago. University of Washington, Seattle, ix + 282 pp. 1974. Keys to the marine invertebrates of Puget Sound, the San Juan Archi- pelago, and adjacent regions. University of Washington, Seattle, x + 226 pp. Lemos de Castro, A. 1965. Crustdceos isé6podos epicarideos do Brasil. IV. Sobre a ocorréncia de Pseudione upogebiae Hay no litoral nordestino (Isopoda, Bopyri- dae). Arq. Est. Biol. Mar. Univ. Ceara 5: 11-14. Lockington, W. N. 1877. Description of a new genus and species of decapod crusta- cean. Proc. California Acad. Sci. 7:55—57. 1878. Remarks upon the Thalassinidae and Astacidea of the Pacific coast of North America, with description of a new species. Ann. Mag. Nat. Hist. ser. 5, 2:299-304. MacGinitie, G. E. 1935. Ecological aspects of a California marine estuary. Amer. Midland Nat. 16:629—765. —. 1937. Notes on the natural history of several marine Crustacea. Amer. Mid- land Nat. 18:1031—1037. , and N. MacGinitie. 1949. Natural history of marine animals. McGraw-Hill, New York, xii + 473 pp. , and 1968. Natural history of marine animals. 2nd. ed. McGraw-Hill, New York, xii + 523 pp. Menzies, R. J., and M. A. Miller. 1954. Key to Chelifera and the suborders of the Isopoda. Pages 138-159, in S. F. Light, R. I. Smith, F. A. Pitelka, D. P. Abbott, and F. M. Weesner, eds. Intertidal invertebrates of the central California coast. Rev. ed. University of California Press, Berkeley and Los Angeles. Miller, M. A. 1975. Phylum Arthropoda: Crustacea, Tanaidacea and Isopoda. Pages 277-312, in R. I. Smith and J. T. Carlton, eds. Light’s Manual: Intertidal in- 818 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON vertebrates of the central California coast. University of California Press, Berkeley and Los Angeles. Nierstrasz, H. F., and G. A. Brender a Brandis. 1923. Die Isopoden der Siboga- Expedition. II. Isopoda Genuina. I. Epicaridea. Siboga-Expeditie Monogr. 32b:57-121. Sand 1931. Papers from Dr. Th. Mortensen’s Pacific Expedition 1914-16. LVII. Epicaridea II. Vidensk. Medd. Dansk Naturh. Foren. K¢ben- havn 91:147-226. Pearse, A. S. 1945. Ecology of Upogebia affinis (Say). Ecology 26:303-305. 1947. Observations on the occurrence of certain barnacles and isopods at Beaufort, N.C. Jour. Washington Acad. Sci. 37:325-328. —. 1953. Three parasitic isopods (Bopyridae) from the Carolina coast. Jour. Parasit. 39:619-620. Richardson, H. 1899a. Key to the isopods of the Pacific coast of North America, with descriptions 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 descriptions of twenty-two new species. Ann. Mag. Nat. Hist. ser. 7, 4:157—187, 260-277, 321-338. — —. 1900. Synopses of North American invertebrates. VIII. The Isopoda.— Part II. Asellota, Oniscoidea, Epicaridea. Amer. Nat. 34:295-309. ——. 1904. Contributions to the natural history of the Isopoda. Proc. U.S. Nat. Mus. 27:1-89. —. 1905. A monograph on the isopods of North America. Bull. U.S. Nat. Mus. plein Sb WDAZ yoyo). Sadoglu, P. 1969. Variation in eye degeneration and pigment in some parasitic iso- pods during their life cycle. Pubbl. Staz. Zool. Napoli 37:173-209. Shiino, S. M. 1952. Phylogeny of the family Bopyridae. Ann. Rept. Prefectural Univ. Mie, Sect. 2, Nat. Sci., 1:33-56. —. 1965. Phylogeny of the genera within the family Bopyridae. Bull. Mus. Nat. Hist. Nat., Paris, sér. 2, 37:462—465. Stebbing, T. R. R. 1893. A history of Crustacea: recent Malacostraca. The inter- national scientific series. Kegan Paul, Trench, Triibner, London, xvii + 466 pp. Stimpson, W. 1857. On the Crustacea and Echinodermata of the Pacific shores of North America. I. Boston Jour. Nat. Hist. 6:444-532. Stromberg, J.-O. 1971. Contribution to the embryology of bopyrid isopods with spe- cial reference to Bopyroides, Hemiarthrus, and Pseudione (Isopoda, Epicaridea). Sarsia 47:1-46. PROC. BIOL. SOC. WASH. 90(4), 1977, pp. 819-825 CERATOLANA PAPUAE, A NEW GENUS AND SPECIES OF MANGROVE-BORING CIROLANID ISOPOD FROM PAPUA NEW GUINEA Thomas E. Bowman Abstract.—Bowman, Thomas E., Department of Invertebrate Zoology, Smithsonian Institution, Washington, D.C. 20560.—Cerotolana papuae, n. gen., n. sp., from burrows in mangroves in Papua New Guinea is described. It is distinguished by fusion of the head with pereonite 1 and the prominent “horns” formed by the rostrum and the frontal lamina. The first specimen of the extraordinary cirolanid isopod described herein was sent to me by Dr. Helmut Kthne, Bundesanstalt fiir Materialprifung, Berlin, West Germany, with the request that I identify it for the collector, Miss Suzanne Rayner, Forest Products Research Centre, Boroko, Papua New Guinea. When I reported to Miss Raynor that her isopod belonged to an undescribed genus and species of Cirolanidae, she kindly sent me a second specimen, together with full collection data. Thanks are tendered to Dr. Kiihne and to Miss Rayner for making it possible for me to describe this remarkable isopod, whose appearance is exquisitely shown by Mrs. Carolyn Bartlett Gast in Fig. 1. I am grateful to Dr. Fenner A. Chace, Jr., for his careful review of the manuscript. Cirolanidae Ceratolana, new genus Body long, rather slender, apparently unable to roll into a ball. Eyes present. Rostrum and frontal lamina produced into prominent “horns.” Head and pereonite 1 fused. Pleonites 1-3 reaching lateral margins of pleon; epimera of pleonite 3 covering lateral margins of pleonites 4-5. Peduncles of antennae | and 2 3-merous and 5-merous respectively. Pereo- pod 1 robust; pereopods 2-7 more slender, ambulatory. Rami of pleopods undivided; exopods of pleopods 3-5 with partial suture laterally; endopods without marginal setae; exopods with marginal setae, numbers of setae de- creasing progressively from pleopod 1 to pleopod 5. Etymology.—From the Greek Cerato (= horn) referring to the modified frontal lamina, + (Ciro) lana. Gender feminine. Type-species.—Ceratolana papuae, new species. Ceratolana papuae, new species Figs. 1-4 Material examined—Papua New Guinea, near Marshall Lagoon, leg. Suzanne Rayner, from mangrove: Mouth of Gonema River, LO°05’S, 145 820 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 1. Ceratolana papuae, lateral, dorsal, ventral and oblique anterodorsal views. Drawings by Carolyn Bartlett Gast. 10’E, 9 March 1975, 6 holotype, 25 mm, USNM 169302. Galley Reach (an inlet), 9°05’S, 146°57’E, 19 April 1975, 6 paratype, 22 mm, USNM 169303. Description.—Body slender, about 34x as long as wide; sides nearly parallel, diverging slightly posteriorly, greatest width at pereonite 7. Head short, fused with pereonite 1; posterior margin indicated by raised trans- 821 VOLUME 90, NUMBER 4 39 ; ‘ Pri thy [th b, Palp of left mandible; ce, Maxilliped. a, Left mandible; ratolana papuae, holotype: Picea. Ce Maxilla 1; d, f, axilla 2; e, Endopod of maxilla 2; M 2) iw) bo PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 3. Ceratolana papuae, holotype, left pereopods: a, Pereopod 1; b, Pereopod 2; c, Pereopod 3; d, Pereopod 4; e, Pereopod 5; f, Pereopod 6; g, Pereopod 7; h, Pereopod 7, carpo-propal suture, medial; i, Same, lateral. 823 VOLUME 90, NUMBER 4 ne se ee ee SES q I | ! ' ! ! ! \ \ b-g Holo- =. Fig. 4. Ceratolana papuae: a, Paratype, pereonite 7 and pleon, lateral; type; b-f, Pleopods 1-5; g, Left uropod, ventral. 824 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON verse bar between posterior bases of ocular lobes; dorsum with depression anterior to transverse bar. Eyes small, borne on lateral surfaces of elevated ocular lobes; each lobe with small conical process posteromedial to cornea and large bifid anterior process. Frons produced anteriorly into conspicuous conical rostrum nearly reaching distal margin of 2nd peduncle segment of antenna 2. Frontal lamina produced anteriorly into subcylindrical horn ending in ventral prong flanked by 2 subapical prongs, and 2 shorter dorsal prongs, reaching apices of subapical prongs. Pereonite 1 with longitudinal middorsal groove in anterior half, connect- ing posteriorly to short transverse groove. All pereonites with surface punctae and inconspicuous surface fuzz. Coxae increasing in size poste- riorly, those of pereonites 5-7 with acutely produced posteroventral cor- ners; coxa 7 overlapping pleonites 1-2. Pleonites 1-3 reaching lateral margins of pleon. Pleonite 4 covering lateral margins of pleonite 5; lateral margins of pleonite 4 covered by much expanded pleura of pleonite 3 which are also expanded anteriorly ventral to pleura of pleonite 2. Telson linguiform; margins hirsute, without spines. Dorsum with median longitudinal ridge having 3 main peaks, and anterolateral longitudinal ridge at base of peduncle of each uropod. Antenna | short, reaching midlength of 4th segment of antenna 2 pe- duncle; flagellum 10-11-merous. Antenna 2 robust, reaching slightly be- yond posterior margin of pereonite 4, flagellum 28-32-merous. Mandibles with 3-cusped incisors; cusps more deeply separated in right incisor. Lacinia of left molar with 7 spines; right lacinia not examined. Segment 2 of palp nearly 3x length of segment 1, with about 21 setae on distal 7; segment 3 falcate, slightly more than % length of segment 2, armed with 26 setae increasing in length distally. Exopod of maxilla 1 with 11 spines and 1 setae between spines at midlength of gnathal surface. Endopod with 3 terminal spines, each with a circle of spinules at mid- length; and a subterminal setule. Maxilla 2 with 7 large spines each on palp and exopod, and 6 setae on exopod; endopod with 18 setae of varying lengths. Maxilliped with 4 retinaculae; palp segments expanded medially, with rather dense setation on both margins. Pereopod 1 shorter and stouter than other pereopods; basis quite broad; ischium and merus with transverse rugae; carpus with incisure on posterior margin; dactyl strongly developed. Pereopods 2-4 successively shorter, moderately slender, posterior margins pubescent, setation very sparse. Pereopods 5-7, distolateral margins of ischium, carpus, and merus armed with robust, blunt spines; distomedial margins with more slender barbed spines; pereopods 5 and 6 subequal in length, slightly shorter than pereo- Powe Appendix masculina of pleopod 2 inserted at proximal % of endopod, reaching distal %4; straight; apex obtuse. VOLUME 90, NUMBER 4 825 Uropod peduncle acutely produced medially. Exopod oblong, much narrower and distinctly shorter than spatulate endopod. Margins of both rami pubescent, armed with setae but without spines. Relationships.—Fusion of the head with pereonite 1 is found in the highly modified Gnathiidea and in certain Epicaridea, but not elsewhere among the Isopoda, as far as I know. The extraordinary modification of the rostrum and especially of the frontal lamina is unique to Ceratolana. The pattern of pleonal segmentation is shared by Creaseriella and Speocirolana (Bowman, 1975), but in neither of these genera are the epimera of pleonite 3 so expanded. The mouthparts of Ceratolana are typically cirolanid, but the relationships of the new genus to other genera of Cirolanidae are not known. Habitat —According to Miss Rayner, Ceratolana was found only in man- groves, in cylindrical burrows up to 25 mm in length and at right angles to the surface of the mangrove. The burrows were in the intertidal region in a part of the estuary in which the salinity could vary from 30% to 5% during a tidal cycle. Day water temperatures here vary from 26°-32°C during the year. It is not known how Ceratolana digs its burrows, but it seems possible that the horns of the rostrum and frontal lamina may be used to rasp the wood. The fusion of the head and pereonite 1 would improve the leverage for rasping movements. Literature Cited Bowman, Thomas E. 1975. A new genus and species of troglobitic cirolanid isopod from San Luis Potosi, Mexico. Occasional Papers Museum Texas Tech University Do d—=7, PROC. BIOL. SOC. WASH. 90(4), 1977, pp. 826-828 DISTRIBUTIONAL NOTES ON NOTIOSOREX AND MEGASOREX IN WESTERN MEXICO Robert D. Fisher and Michael A. Bogan The known distributions of Notiosorex crawfordi and Megasorex gigas were summarized recently by Armstrong and Jones (1972a and 1972b, respectively). We here present additional distribution records for these taxa. Curatorial work on North American soricids in the National Museum of Natural History (USNM) has resulted in the discovery of the only known specimen of N. crawfordi from the state of Zacatecas, Mexico. The speci- men, an adult female (USNM 90845), was collected by E. W. Nelson and E. A. Goldman on 4 September 1897 near Plateado, Zacatecas. According to Goldman (1951:288), Plateado is a small village near the northern end of Sierra Moroni, 28 mi SE of Colotlan, Jalisco. Our map (American Geo- graphical Society, NF13) shows the airline distance between the two places to be 24 km. Nelson and Goldman worked around Plateado and the ad- jacent slopes of Sierra Moroni from 31 August to 4 September 1897 and all specimens collected in this vicinity were labelled “Plateado.” Nelson's report (USNM archives) on this area states that the Notiosorex and two specimens of Sorex saussurei (USNM 90844 and 90846) were taken “in damp spots under rocky ledges in the Sierra Moroni” and were “not com- mon. The precise elevation at which the Notiosorex was captured is un- known but was at least 7,600 ft (the elevation of Plateado) and may have been as high as 8,500 ft, the highest elevation given for Sierra Moroni. Either of these figures increases the known elevational range of this species from the previously reported 7,000 ft in Arizona (Lange, 1959). External measurements (in mm) as given on the label are: total length, 94; tail, 28; and hind foot, 12. Although the specific identity can be ascertained from the skull, the skull is too badly fragmented to measure. We tentatively assign the specimen to N. c. evotis on the basis of the external measure- ments given by Armstrong and Jones (1972a) for N. crawfordi. Twente and Baker (1951) identified four specimens from near Guadalajara, Jalisco, only as N. crawfordi and noted that N. c. evotis was then known only from Mazatlan, Sinaloa. Baker and Alcorn (1953) assigned three specimens from Michoacan to N. c. evotis on geographic grounds. Armstrong and Jones (1972a) assigned all the Jalisco and Michoacan material to N. c. crawfordi. Resolution of subspecific relationships among Notiosorex from this area of México must await additional comparative material. Schlitter (1973) first reported N. c. evotis from Nayarit (San Cayetano, VOLUME 90, NUMBER 4 827 5 mi W Tepic). We are now aware of two additional Nayarit specimens of this shrew: an adult female (NAU 1904, skeleton only), taken 15 June 1971 at 1 mi S Compostela; and another adult female (USNM 508358) captured on 11 March 1975 near El Refilion. External and cranial measurements of these two are as follows (Compostela specimen first; cranial measurements after Jackson, 1928): total length, 100, 99; tail, 34, 30; hind foot, 13, 13; ear, —, 8; condylobasal length, 17.4, 18.8; cranial breadth, 8.1, 8.5; palatal length, 7.2, 7.7; least interorbital breadth, 4.5, 4.1; maxillary breadth, 5.4, 5.5; max- illary toothrow, 6.0, 6.5. These measurements are close to those for animals from southern Sinaloa (Armstrong and Jones, 1972a), although greater values in some dimensions may indicate a cline with southern animals being larger. It is of interest that recent field work in Nayarit has not yielded any specimens from the northern part of that state. The southern- most record in Sinaloa is near Escuinapa (Baker, 1962). Nonetheless, we expect N. c. evotis to occur at low to moderate elevations throughout Nayarit. Megasorex gigas was first reported from Nayarit from 7.3 mi ESE Amat- lan de Canias, 5,000 ft (Jones, 1966). This specimen was taken in a moist situation among lava rocks. Two Nayarit specimens of this relatively rare shrew are in the collection of The Museum, Michigan State University (MSU). One, an adult male (MSU 17086), was taken 16 July 1971 at Mineral de Tigre, 6 mi E Huajicori, 1,340 ft; the other, an adult female (MSU 16102), was captured 9 mi WSW Compostela, 2,000 ft on 16 July 1970. On 31 October 1975, A. L. Gardner found a dead adult male Mega- sorex (USNM 511264) floating in a small water tank 3 mi N Coapan, Nayarit. The animal, which was found at 1000 hr was in fresh condition and must have fallen into the tank earlier in the morning. This locality is also characterized by an abundance of lava rocks but the habitat is a xeric one, disturbed by grazing. Measurements of these three specimens (in the order listed above) are: total length, 118, 124, 121; tail, 45, 43, 40; hind foot, 15, 16, 16; ear, 9, 12, 9; condylobasal length, —, 21.2, 22.0; cranial breadth, —, 10.1, 10.8; palatal length, 9.9, 9.5, 9.8; interorbital breadth, 5.1, 5.1, 5.2; maxillary breadth, 6.4, 6.4, 6.7; maxillary toothrow, —, 7.9, 8.6. Values for these measurements are slightly smaller than those given by Armstrong and Jones (1972b) for a small series of M. gigas from Guerrero. The specimen from near Huajicori extends the known range of M. gigas approximately 220 km northward from Amatlan and would indicate that the giant shrew occurs throughout the state at low elevations. We thank Rollin Baker, The Museum, Michigan State University, and Terry Vaughn, Museum of Vertebrates, Northern Arizona University, for the loan of specimens for examination and inclusion in this report. Don 828 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Wilson and Al Gardner read the manuscript. Senor Mario Luis Cossio G., Director de la Direccidén General de la Fauna Silvestre kindly granted the collecting permits which facilitated our work in Nayarit. Literature Cited Armstrong, D. M., and J. K. Jones, Jr. 1972a. Notiosorex crawfordi. Amer. Soc. Mamm., Mammalian Species 17:1-5. , and 1972b. Megasorex gigas. Amer. Soc. Mamm., Mammalian Spe- cies 16:1-2. Baker, R. H. 1962. Additional records of Notiosorex crawfordi from Mexico. Jour. Mamm. 43:283. , and A. A. Alcom. 1953. Shrews from Michoacan, Mexico, found in barn owl pellets. Jour. Mamm. 34:116. Goldman, E. A. 1951. Biological investigations in Mexico. Smithsonian Misc. Coll. 115:1-476. Jackson, H. H. T. 1928. A taxonomic review of the American long-tailed shrews (genera Sorex and Microsorex). N. Amer. Fauna 51:1—-238. Jones, J. K., Jr. 1966. Recent records of the shrew, Megasorex gigas (Merriam), from western Mexico. Amer. Midland Nat. 75:249-250. Lange, K. I. 1959. Soricidae of Arizona. Amer. Midland Nat. 61:96-108. Schlitter, D. A. 1973. Notiosorex crawfordi evotis from Nayarit. Southwest. Nat. 17: 423. Twente, J. W., and R. H. Baker. 1951. New records of mammals from Jalisco, Mex- ico, from barn owl pellets. Jour. Mamm. 32:120-121. Fisher, Robert D., and Michael A. Bogan, National Fish and Wildlife Laboratory, National Museum of Natural History, Washington, D.C. 20560 PROC. BIOL. SOC. WASH. 90(4), 1977, pp. 829-830 ON THE GENERA ECHINASTER MUELLER AND TROSCHEL AND OTHILIA GRAY, AND THE VALIDITY OF VERRILLASTER DOWNEY (ECHINODERMATA: ASTEROIDEA) Enrico Tortonese and Maureen E. Downey Abstract.—Tortonese, E., Instituto Zooprofilattico, Genoa, Italy; and M. E. Downey, Department of Invertebrate Zoology, Smithsonian Insti- tution, Washington, D.C. 20560.—An important taxonomic character, i.e., glassy tubercles on the principal plates, is discussed. The genus Verrillaster Downey is synonymized with Echinaster Mueller and Troschel, and the genus Othilia Gray is revived to accommodate those species of Echinaster in which glassy tubercles are present. The genus Echinaster Mueller and Troschel (1840) includes a number of well-known seastars, distributed mainly in tropical and subtropical shallow waters of the world. The genus is still in need of an adequate definition, as are several of the species, chiefly those occurring in the Western Atlantic. Patches of glassy tubercles occur on many principal plates of some of the species assigned to Echinaster. The patches are variable in size, but as far as we know, size differences are interspecific, no intraspecific variations having been observed. Such patches seem, therefore, to be a good taxonomic character, even if they are not always easily detected on _ poorly preserved specimens. The presence or absence of glassy tubercles has not been determined in all species. One of us (Downey, 1973) found them on most of the species of Echinaster from the Western Atlantic (E. serpentarius, E. echinophorus, E. modestus, etc.) and regarded them as a diagnostic feature of the genus. The same author consequently separated E. spinosus Verrill, which lacks patches of glassy tubercles, from the other _ Western Atlantic species, and erected a new genus, Verrillaster, for this _ species and any others that lack glassy tubercles. One of us (Tortonese) has found that the separation of Echinaster and Verrillaster on such grounds is unacceptable, since glassy tubercles are absent from the type-species of Echinaster, Asterias seposita Retzius (see Mortensen, 1925, re the name seposita and its authorship). Caretul ex- amination of a number of specimens of E. sepositus of various sizes from several locations in the Mediterranean and the Eastern Atlantic (Roscoft and Senegal) revealed that no patches of glassy tubercles were present. Verrillaster (type-species E. spinulosus Verrill) thus becomes a junior synonym of Echinaster, as in both genera the type-species is devoid of glassy tubercles. 830 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON In general appearance, E. sepositus is rather similar to two Indo-West Pacific species, E. purpureus (Gray) and E. luzonicus (Gray), all three species having short spines. Examination has now shown that these Indo- West Pacific species also lack glassy tubercles. This condition supports Tortonese’s (1954) suggestion that E. sepositus is more closely related to E. purpureus from the Red Sea and the Indian Ocean (and also to E. luzonicus from farther east) than to the Western Atlantic species. The more tropical fauna of the latter includes a complex of species or “forms” sharply contrasting with the single one (E. sepositus) found in the less tropical Eastern Atlantic. For the genus comprising the species having glassy tubercles, we pro- pose the oldest available name, Othilia Gray (1840) (type-species Asterias echinophora Lamarck: see Fisher, 1913). The genus Othilia includes the species serpentaria, echinophora, senta, modesta, and brasiliensis, all from the Western Atlantic, and possibly others. It would be interesting to know if and how the presence of patches of glassy tubercles is correlated with other morphological features. If there is a correlation, the recognition of two genera would appear still more acceptable, but a revision of Echinaster on a world-wide basis is, of course, required. Literature Cited Downey, M. E. 1973. Starfishes from the Caribbean and the Gulf of Mexico. Smithsonian Contrib. Zool. No. 126, vi + 158 pp., 2 figs., 48 pls. Fisher, W. K. 1913. A new species of Echinaster, with a note on the name Othilia. Zool. Anz. Leipzig 42:193-196. Mortensen, Th. 1925. The name Echinaster sepositus (Retzius), with remarks on Retzius’ “Dissertatio.” Ann. Mag. Nat. Hist. London, Ser. 9, 16: 546-547. Tortonese, E. 1954. Zoogeografia e speciazione nel gen. Echinaster (Asteroidi). Boll. Zool. 21(2):419-428, 2 figs. PROC. BIOL. SOC. WASH. 90(4), 1977, pp. 831-838 NOTES ON A COLLECTION OF CRABS (CRUSTACEA: BRACHYURA) FROM THE EAST COAST OF MEXICO Jack A. Rickner Abstract.—Rickner, J. A., Department of Wildlife and Fisheries Sciences, Texas A&M University, College Station, Texas 77843.—Representatives of six families of Brachyura are reported as range extensions from the poorly known east coast of Mexico north of Veracruz: Dromiidae, Calappidae, Leucosiidae, Grapsidae, Gecarcinidae, Ocypodidae. This report deals with six families of brachyuran crabs collected from the poorly known east coast of Mexico between Anton Lizardo, Veracruz and the Rio Grande River. The crabs were collected during visits to two coral reefs, Lobos and Enmedio, and a lava outcrop, Punto del Morro, during the summer of 1973. Also included in this report are small amounts of _ material from other sources that provide additional locality data for a few species. Description of Study Areas Two near shore emergent coral reefs were extensively collected (May- June 1973), and limited sampling was made of several other locations. Lobos Reef (21°28’N, 97°13’W), located 120 km southeast of Tampico, is one of a cluster of 3 reefs which comprises the northernmost emergent coral reefs in the western Gulf. It is situated near the boundary between 2 zoogeographical provinces, tropical to the south and warm temperate to the north (Briggs 1974:218-221). Enmedio Reef (19°06’N, 95°56’W), located _ 24 km southeast of Veracruz, is associated with the Veracruz-Anton Lizardo Reef complex. Both reefs have shallow water lagoons and small vege- _ tated islands. The upper 4 m of the reef and the lagoon area were ex- amined during 9 days of collecting on each island. Other areas along the coast that were visited for short periods were Punto del Morro and Tamiahua in the state of Vera Cruz and La Pesca, Tamaulipas. El Morro is an area of lava cliffs projecting into the otherwise smooth continuous sand beach 35 km northwest of Veracruz. Le Pesca and Tamiahua are small fishing villages located on coastal bays north and south, respectively, of Tampico. Material and Methods Collecting on Lobos and Enmedio was made using SCUBA, mask and snorkel, or by wading. Other areas were collected while walking along the shore line. 832 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON | Papers relied upon for identification and distribution data were Chace and Hobbs (1969), Felder (1973), Holthuis (1958, 1959), and Rathbun } (1918, 1933, 1937). Representative specimens are placed in the Texas A&l | University Crustacean Collection (TAIC). Account of Species This report considers 6 families, 17 genera, and 19 species. Many of these species are new locality reports and a few are range extensions into the western Gulf of Mexico. These range extensions reflect the general lack of collecting in this region rather than the geographical advancement into new areas. Abbreviations used for carapace length and width measure- ments are cl and cw. Ovigerous females (ov) are noted. Dromiidae Dromidia antillensis Stimpson 1859 Material examined.—TAIC-432, Enmedio, 1¢é; Author’s collection, El Morro, 1¢é. Measurements—Males: cl 14.1 and 22.5 mm; cw 15.1 and 21.9 mm. Range.—North Carolina to Brazil and West Indies, Surinam (Holthuis 1959), Texas and Yucatan (Rathbun 1937). Remarks.—The specimen from Enmedio had a compound ascidian at- tached to its carapace. Holthuis (1959) reported both individuals collected from Surinam carrying compound ascidians. Hildebrand (1955) reported these crabs to be common on the Campeche Bank wherever ascidians are found. The individual taken at Enmedio was taken from the pier area in water less than 2 m in depth. Dromia erythropus (George Edwards 1771) Material examined.—T AIC-435, Enmedio, 12. Measurements.—Female: cl 70.8 mm, cw 87.4 mm. Range.—Florida to Brazil, Bermuda (Rathbun 1937), Texas and Lou- isiana (Felder 1973). Lobos and Enmedio (Ray 1974). Remarks.—This individual was collected from under debris near the pier area of Enmedio in water less than 2 m deep. Although this species is known to carry a covering of sponge or ascidians, neither was found on the carapace of this individual. Calappidae Clappa flammea (Herbst 1794) Material examined.—TAIC-414, Enmedio, 1¢é; TAIC-416, Enmedio, lice Measurements.—Males: cl 27.7 and 29.3 mm, cw 37.5 and 39.7 mm. VOLUME 90, NUMBER 4 833 Range.—North Carolina to the Florida Keys, Bermuda to the Bahamas, the Tortugas and the Gulf Coast of the United States and Mexico (Holthuis 1958). Remarks.—Both individuals were collected at the pier on Enmedio in water depth of less than 2 m. One individual was taken from under debris, and the other was collected while being eaten by a small octopus. This species has been reported from the Gulf of Campeche by Holthuis (1958) and Hildebrand (1955). Calappa gallus (Herbst 1803) Material examined.—TAIC-413, Enmedio, 24 ¢. Measurements.—Males: cl 35.0 and 35.6 mm, cw 46.1 and 48.8 mm. Range.—Florida Keys to Brazil, Bermuda, West Africa, Red Sea, Persian Gulf, islands of the Indian Ocean and Pacific Ocean, and Mexico (Rathbun 1937). Remarks.—Both individuals were collected during night dives in the shallow lagoon, west of Isla de Medio. Water depth was less than 2m. Chavez et al. (1970) reported this species as occurring on Lobos Reef. Hepatus epheliticus (Linnaeus 1763) Remarks.—One individual was seen floating in the boat channel at Lobos Island. It may have been brought into the area by one of the many shrimp trawlers that frequent the island. Leucosiidae Ebalia cariosa (Stimpson 1860) Material examined.—TAIC-437, Enmedio, 14, 12. Measurements.—Male: cl 9.9 mm, cw 10.2 mm; female: cl 7.4 mm, cw o.1 mm. Range.—North Carolina to Florida, Jamaica, and Brazil (Rathbun 1937) and Surinam (Chace, personal communication). Remarks.—This appears to be the westernmost record for this species, there being no previous record from the east coast of Mexico. It is known to occur from below the low tide line to 45 m. These individuals were col- lected at night in the lagoon area west of the island in water less than 2 m in depth on a bare coral sand bottom. Grapsidae Platychirograpsus typicus Rathbun 1914 Material examined.—Author’s Collection, Rio Colipa, 2¢ 6, 1°, 834 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON | Measurements.—Males: cl 29.3 and 29.5 mm, cw 34.3 and 34.7 mm; female: cl 27.0 mm, cw 31.5 mm. Range.—Rio Macuspana, Tabasco, Mexico to the Rio de Tecolutla, Vera- cruz, Mexico (Pieltain 1945). Remarks.—These individuals were collected from the Rio Colipa, 20 miles south of Nautla, Veracruz. Rathbun (1918) reported this species as occurring 140 miles from sea at an altitude of over 100 feet on the Macu- spana River. Percnon gibbesi (Milne Edwards 1853) Material examined.—TAIC-322, Enmedio, 12 (ov); TAIC-325, Lobos, Ig, Le (Oy). Measurements.—Male: cl 16.9 mm, cw 14.5 mm; females (ov): cl 17.4 and 26.1 mm, cw 16.2 and 24.4 mm. Range.—Florida and Bahamas to Brazil, Bermudas, and Clarion Island, Mexico (Rathbun 1918). Remarks.—Individuals taken at Lobos Island were collected from around the concrete well platform in the lagoon. The one individual collected at Enmedio was taken from the windward reef crest where coral boulders are exposed during low tide. As far as I have been able to determine, this is the first report of Percnon gibbesi from the east coast of Mexico. Goniopsis cruentata (Latreille 1803) Material examined.—T AIC-487, Tamiahua, 1°. Measurements——Female: cl 29.5 mm, cw 35.9 mm. Range.—East coast of America, Bermuda, Bahamas, West Indies to Brazil and the west coast of Africa, Surinam (Holthuis 1959) and Texas (Felder 1973). Remarks.—This individual was collected from a mangrove area near Tamiahua, Veracruz. Grapsus grapsus (Linnaeus 1758) Material examined.—T AIC-282, El Morro, 2é ¢; TAIC-290, El Morro, 12; TAIC-286, Enmedio, 1¢; TAIC-357, Enmedio, 22 2. Measurements.—Males: cl 43.7-54.5 mm, cw 49.2-61.5 mm; females: cl 20.4-38.8 mm, cw 23.3-45.7 mm. Range.—Florida, Bermudas, West Indies, to Brazil, the eastern Atlantic from southern Portugal to northern Angola, the eastern Pacific from cen- tral Baja California to central Chile (Chace and Hobbs 1969), Port Aransas, Texas (Felder 1973), and Lobos Island (Chavez et al. 1970). Remarks.—Three individuals were collected from the pier on Enmedio. They were not abundant probably due to the lack of hard substrate per- VOLUME 90, NUMBER 4 835 manently above the tide line. Numerous individuals were seen moving over the lava cliffs at El Morro. Although none was taken at Lobos Island, they were occasionally seen along a coral rubble ridge adjacent to the boat channel. Individuals were also seen on the rock jetties at Tuxpan and Veracruz. Cyclograpsus integer Milne Edwards 1837 Material examined.—TAIC-415, Lobos, 22 2; TAIC-467, El Morro, 34 4, IL2 (Ow). Measurements.—Males: cl 6.2-8.5 mm, cw 7.7-10.6 mm; females (non-ov): cl 5.3 and 10.6 mm, cw 6.8 and 14.4 mm; female (ov): cl 8.2 mm, cw 10.5 mm. Range.—Bermuda, Florida to Brazil, eastern Atlantic from Senegal to Congo (Chace and Hobbs 1969), Texas (Felder 1973), western Indian Ocean at Madagascar, Indonesia, northern west Pacific, and eastern cen- tral Pacific (Griffin 1968). Remarks.—The individuals from Lobos Reef were taken from the splash zone of the coral rubble spoil islands along the boat channel and at the splash zone of the lava cliffs of El Morro. Geograpsus lividus H. Milne Edwards 1837 Material examined.—TAIC-287, Lobos, 644; TAIC-297, Lobos, 1°. Measurements.—Males: cl 23.6-30.8 mm, cw 26.9-36.4 mm; female: cl 20.8 mm, cw 25.7 mm. Range.—Bermudas, Florida Keys, West Indies to Brazil, eastern At- lantic from Senegal to northern Angola, eastern Pacific from southern Baja California to northern Chile, and Hawaii (Chace and Hobbs 1969). Remarks.—These individuals were collected from under coral rubble at the tide line on the spoil islands of Lobos. Sesarma cinereum (Bosc 1801 or 1802) Material examined.—Author’s collection, Lobos, 346 ¢, 12 (ov). Measurements.—Males: cl 12.2-13.3 mm, cw 13.8-14.2 mm; female (ov): cl 14.3 mm, cw 15.8 mm. Range.—Chesapeake Bay, Md. to British Honduras, West Indies to Venezuela (Williams 1965). Abundantly found throughout the northwestern Gulf of Mexico (Felder 1973). Remarks.—All individuals were taken from coral rubble near or at the water line. Specimens collected from the Rio Colipa were also examined. Pachygrapsus gracilis (Saussure 1858) Material examined.—TAIC-412, Lobos, 2¢6 6, 222 (non-ov), 12 (ov). Measurements.—Males: cl 5.1 and 5.9 mm, ew 6.9 and 7.5 mm: females 836 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON (non-ov): cl 5.5 and 6.7 mm, cw 7.4 and 8.8 mm; females (ov): cl 8.0 mm, cw 10.0 mm. Range.—Bermudas, Florida, West Indies to Brazil; eastern Atlantic from Senegal to the Congo (Chace and Hobbs 1969); Texas (Felder 1973). Remarks.—Individuals were collected at night from the spoil islands and coral rubble along the boat channel at Lobos. Pachygrapsus transversus (Gibbes 1850) Material examined.—TAIC-409, Lobos, 74 6, 42 2 (non-ov), 1222 (ov). Measurements——Males: cl 6.7-14.4 mm, cw 8.6-18.7 mm; females (non- ov): cl 5.8-11.6 mm, cw 7.9-14.8 mm; females (ov): cl 6.3-11.7 mm, cw 8.6- 15.9 mm. Range.—Bermudas, North Carolina to the West Indies and Uruguay; east- ern Atlantic from the Mediterranean to northern Angola; eastern Pacific from California to Peru (Chace and Hobbs 1969); Louisiana (Behre 1950); Texas (Whitten, Rosene, and Hedgpeth 1950); and Mexico (Chavez et al. 1970). Remarks.—These individuals were collected from the coral rubble areas of Lobos and from the lava cliffs of El Morro. This species was found abundantly at the intertidal zone. Gecarcinidae Cardisoma guanhumi Latreille 1825 Material examined.—TAIC-485, La Pesca, 16; TAIC-486, La Pesca, Ae Measurements.—Males: cl 47 and 52.2 mm, cw 59.4 and 64.7 mm. Range.—Bermuda, Florida, the Bahamas to Brazil and the West Indies (Holthuis 1959), Louisiana (Behre 1950), and Texas (Rathbun 1918). Remarks.—These individuals were taken from their terrestrial burrows along clay banks at La Pesca. Numerous other individuals were seen at the entrances to their burrows. Gecarcinus lateralis (Freminville 1835) Material examined.—TAIC-278, Enmedio, 24 ¢; TAIC-279, El Morro, 12 (ov); TAIC-280, El Morro, 16; TAIC-284, Lobos, 24 3, 32 2: TAIC-296, Lobos, 2¢ 8, 12 (ov); TAIC-333, El Morro, 22 é. Measurements.—Males: cl 29.2-48.1 mm, cw 29.5-33.8 mm: females (non-oy): cl 24.4-29.1 mm, cw 29.5-33.8 mm; females (ov): cl 31.0 and 43-2 MM, CW of.o and 53.2 mm. Range.—Bermudas and southern Florida to French Guiana (Chace and Hobbs 1969), Texas (Ray 1967), and Lobos Island, (Chavez et al. 1970). Remarks.—This land crab was found abundantly well above the tide VOLUME 90, NUMBER 4 837 line. At El Morro individuals were found in plowed fields some distances from the beach. Although individuals were not collected at Tuxpan, La Pesca, Anton Lizardo, and Veracruz, they were seen along the beach. Several small individuals were examined from a mangrove area near Tamiahua. Ocypodidae Ocypode quadrata (Fabricius 1787) Material examined. —TAIC-281, El Morro, 24 ¢; TAIC-293, Lobos, 14; TAIC-300, Enmedio, 12; TAIC-301, Lobos, 2¢ 6, 42 ?; TAIC-324, Lobos 1é. Measurements.—Males: cl 12.1-36.4 mm, cw 14.9-43.9 mm; females: cl 15.7-26.5 mm, cw 19.8-33.0 mm. Range.—Rhode Island to Florida, West Indies, Bermuda, Brazil (Chace and Hobbs 1969), Louisiana (Behre 1950), Texas (Hedgpeth 1953), and Mexico (Chavez et al. 1970). Remarks.—These were common burrowers on the sandy beaches above the high tide mark. They were seen abundantly at night, late after- noon, and early morning. I found these crabs in abundance on the beaches from the Rio Grande to Anton Lizardo. Ucides cordatus (Linnaeus 1763) Material examined.—TAIC-441 and 442, Tamiahua, 26 ¢. Measurements.—Males: cl 42.4 and 42.9 mm, cw 55.9 and 56.7 mm. Range.—Florida, West Indies, and Panama to Southeastern Brazil (Holthuis 1959, Chace and Hobbs 1969). Remarks.—These individuals were collected from a grove of mangroves along the banks of Laguna de Tamiahua, Mexico. Chace and Hobbs (1969) transferred Ucides from the family Gecarcinidae to the family Ocypodidae. This species has not been reported previously from Mexico. Acknowledgments I would like to express my appreciation to Dr. J. W. Tunnell and Dr. A. H. Chaney who made possible these collecting trips. I am grateful to Dr. Ned Strenth and Mr. and Mrs. Steve Rabalais who made available speci- mens which they collected at Rio Colipa and Tamiahua. To Dr. F. A. Chace, Jr., I am indebted for identification of a doubtful specimen. I also wish to express my thanks to Dr. Chaney for verification of specimens. Literature Cited Behre, E. H. 1950. Annotated list of the fauna of the Grand Isle region. Occas. Papers Mar. Lab. Louisiana State Univ. 6:1-66. 838 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Briggs, J. C. 1974. Marine zoogeography. McGraw-Hill Book Co., New York, 475 pp. Chace, F. A., and H. H. Hobbs. 1969. The freshwater and terrestrial decapod crusta- ceans of the West Indies with special reference to Dominica. Bull. U.S. Nat. Mus. 292: 258 pp., figs. 1-76. Chavez, E. A., E. Hidalgo, and M. L. Sevilla. 1970. Datos acerca de las comunidades benténicas del Arrecife de Lobos, Veracruz. Rev. Soc. Mexicana Hist. Nat. 31: 211-280, pls. 15, figs. 1-96. Felder, D. L. 1973. An annotated key to crabs and lobsters (Decapoda, Reptantia) from coastal waters of the northwestern Gulf of Mexico. Cent. Wetland Resources, Louisiana State Univ. Publ. No. LSU-SG-73-02. 103 pp. pls. 1-12. Griffin, D. J. G. 1968. A new species of Cyclograpsus (Decapoda, Grapsidae) and notes on five others from the Pacific Ocean. Crustaceana 15(3):235-248. Hedgpeth, J. W. 1953. An introduction to the zoogeography of the northwestern Gulf of Mexico with reference to the invertebrate fauna. Publ. Inst. Mar. Sci. Univ. Texas 3(1):107-224, 46 figs. Hildebrand, H. H. 1955. A study of the fauna of the pink shrimp grounds in the western Gulf of Campeche. Publ. Inst. Mar. Sci. Univ. Texas 4(1):170-232, tabls. 1-7. Holthuis, L. B. 1958. West Indian crabs of the genus Calappa, with a description of three new species. Studies Fauna Curacao 8(7):146-186, 27 figs. —. 1959. The Crustacea Decapoda of Suriname (Dutch Guiana). Zool. Verh. Leiden 44:1-296, 68 text-figs., 16 pls., 1 map. Pieltain, C. B. 1945. Notas sobre Platychirograpsus typicus. Ciencia 6:267—270. Rathbun, M. J. 1918. The grapsoid crabs of America. Bull. U.S. Nat. Mus. 97:461 pp. 1933. Brachyuran crabs of Porto Rico and the Virgin Islands. Scientific Survey of Porto Rico and the Virgin Islands, New York Academy of Sciences 15(1):1-121, figs. 1-107. — —. 1937. The oxystomatous and allied crabs of America. Bull. U.S. Nat. Mus. 166:278 pp. Ray, C. 1967. Gecarcinus lateralis Freminville in Texas. Texas Jour. Sci. 19(1):109. Ray, J. P. 1974. A study of the coral reef crustaceans (Decapoda and Stomatopoda) of two Gulf of Mexico reef systems: West Flower Garden, Texas and Isla de Lobos, Veracruz, Mexico. Ph.D. Diss., Texas A&M Univ. 322 p. Whitten, H. L., H. F. Rosene, and J. W. Hedgpeth. 1950. Invertebrate fauna of the Texas coast jetties. Publ. Inst. Mar. Sci. Univ. Texas 1(2):53-87. Williams, A. B. 1965. The decapod crustaceans of the Carolinas. U.S. Fish. Bull. 65(1'): 1-298, figs. 1-252. PROC. BIOL. SOC. WASH. 90(4), 1977, pp. 839-848 ANTHURIDS FROM THE WEST COAST OF NORTH AMERICA, INCLUDING A NEW SPECIES AND THREE NEW GENERA (CRUSTACEA, ISOPODA) George A. Schultz Abstract.—Schultz, George A., 15 Smith St., Hampton, New Jersey 08827. —A review is given of the seven named and two unnamed species of anthurid isopods recorded from the west coast of North America. Two new genera, Califanthura and Silophasma are proposed respectively for Colanthura squamosissima Menzies, 1951, and Haliophasma geminata Menzies and Barnard, 1959. A new genus and species, Cortezura penascoensis, from the Gulf of California, is described. _ The anthurids of the west coast of North America include 7 named species, each in a separate genus. The citation by Brusca (1973) of Mesanthura sp. _ adds another if that species proves not to be M. occidentalis Menzies and Barnard (1959). Also Menzies (1962) added an “Anthurid” which was not more exactly named. With the new species described herein 8 named spe- cies of anthurids and 2 anthurids of uncertain status are recorded from _ the west coast of North America. One of the established genera is abolished and one is redefined so as to exclude the species from the west coast, so 2 new genera are proposed for established species. One new genus is _ described here for a new species so that the total number of named genera of anthurids from the west coast of North America, 8, also equals the number of species present. A key to the 8 named species follows the dis- cussions of the individual species. Califanthura, new genus - Colanthura Richardson, 1902.—Menzies, 1951:15.—Menzies and Barmard, . 1959:15. nec Colanthura Richardson, 1902.—Schultz, 1969:89. The genus Colanthura Richardson (1902) is based on C. tenuis Richard- son (1902) (type-species by monotypy) from Bermuda. Colanthura tenuis, as described by Richardson, is based on 2 tiny specimens each with 6 large peraeonal segments and a pleon composed of very short segments. The specimens are identical with tiny specimens of Paranthura infundib- ulata Richardson (1902), which also have a short pleon composed of very short segments (personal observation and comparison of type-specimen and other specimens in Peabody Museum, Yale University). Since the type- 840 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON species of the genus is not a valid species, the genus is invalid and the generic name must be supressed. Menzies (1951) placed the species squamosissima in Colanthura Richard- son so that it must now be placed in another genus or a new genus must be created to include it. Menzies (1951, p. 15) used the generic name without discussion. Menzies and Barnard (1959) gave a diagnosis of the genus based mostly on C. squamosissima. The diagnosis of the new genus Califanthura follows that of Menzies and Barnard (1959, p. 15) with some modifications. | Diagnosis.—Eyes small. Mouthparts for piercing and sucking. Peraeonal — segment VII vestigal. Pleon short with distinct sutures. Telson flattened, not indurate, without paired statocysts. Maxillipedal palp triarticulate. Palm of peraeopod I smooth, with prominent basal tooth. Peraeopods I[V-— VI with carpus not underwriting propodus. Pleopods 1 not indurate. Etymology and gender.—The generic name Califanthura is formed from a combination of California and the suffix “-anthura.”. The gender is feminine. Type-species.—Colanthura squamosissima Menzies, 1951. Califanthura squamosissima (Menzies) Colanthura squamosissima Menzies, 1951:115, figs. 14-16—Menzies and Barnard, 1959:15, fig. 9A, B.—Schultz, 1969:90, 117. The species is distinguished from other anthurids on the west coast be- cause it lacks a well developed peraeonal segment VII. What appears to be a vestige of that segment appears in the illustration of Menzies (1951, p. 115, fig. 14a) just in front of the pleon. Menzies recorded a gravid female 5.2 mm long. The species is well described and illustrated by Menzies (1951) and also diagnosed by Menzies and Barnard (1959). It is found from Marin County, California, south to the Mexican border in depths from 18.3-90.1 m. Silophasma, new genus Haliophasma Haswell—Menzies and Barnard, 1959, p. 17. The genus Haliophasma Haswell, 1881, was described from species from Australia, and H. purpureum is the type-species (Poore, 1975 p. 503—type by virtual monotypy). Poore restricted the genus to species with partic- ular morphological characters, redescribing the Australian species. Here the species H. geminatum Menzies is placed in a new genus Silophasma since it does not conform to Poore’s new restricted definition of Haliophasma Haswell. Poore excludes H. geminatum from the genus because it lacks dor- sal grooves and pits. The diagnosis of Haliophasma by Menzies and Barnard (1959) was based mainly on H. geminatum. They stated originally that it had “piercing VOLUME 90, NUMBER 4 841 and sucking’ mouthparts, but afterwards Menzies (1962) corrected this to chewing mouthparts. In the later work Menzies recorded immature specimens of what he tentatively identified as the species. The species is here placed in the new genus Silophasma which is diagnosed as follows: Diagnosis.—Eyes present. Mouthparts for chewing. Statocysts paired. Telson sculptured. No dorsal grooves or pits. Carpus of peraeopods IV- VII not underriding propodus. Pleopods 1 operculate. Maxillipedal palp of 4 articles. Peraeonal segment VII shortest of all. Pleon longer than peraeonal segment VII with sutures indistinct. Male with extremely long antenna 2. Etymology and gender.—‘Silo-” is without meaning and “-phasma,” the suffix of the former generic name Haliophasma, means ghost in Latin. The gender is neuter. Type-species.—Haliophasma geminatum Menzies and Barnard, 1959. Poore (1975) stated that Haliophasma Haswell is neuter, so Menzies and Barnard should have used geminatum, not geminata, as their name for the species. Silophasma geminatum (Menzies and Barnard) Haliophasma geminata Menzies and Barnard, 1959:17, figs. 11-12.—Menzies, 1962:339.—Schultz, 1964:312; 1966:13; 1969:103, fig. 141.—Iverson, 1974: 165. Haliophasma geminatum Menzies and Barnard.—Poore, 1975:531. The species has been recorded from Monterey Bay to Bahia de San Quintin, Baja California, in stations in depths of 9.2-512.4 m. The range also includes stations near Santa Catalina Island. It ranges to 7 mm long, and is apparently quite common on the continental shelf. Cyanthura munda Menzies Cyanthura munda Menzies, 1951:111, figs. 112, 13—Menzies and Barnard, 1959:16, fig. 10A, B.—Schultz, 1964:312; 1969:105, fig. 144a-—c. A female 6 mm long was the largest recorded by Menzies (1951). It has been recorded from Marin County, California, to the Mexican border in depths of from 18.3-55 m. Paranthura elegans Menzies Paranthura elegans Menzies, 1951:106, figs. 9-11; 1962:340.—Menzies and Bamard, 1959:19, fig. 13—Schultz, 1969:94, fig. 125a-e. The species, which ranges to 9.1 mm long, has been collected from Marin County, California, to Bahia de San Quintin, Baja California, from shallow water to a depth of 55 m. 842 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Bathura luna Schultz Bathura luna Schultz, 1966:12, pl. 7; 1969:98, fig. 132. The genus contains a single blind species which ranges to 21 mm long. It was taken from Tanner Canyon on the continental shelf off Los An- geles at depths between 783 and 812 m. It has not been reencountered since its initial description. Apanthura californiensis Schultz Apanthura californiensis Schultz, 1964:312, fig. 4; 1969:99, fig. 135. The species which ranges to 11 mm long was taken off Santa Monica, California, at a depth of 80 m. It has not been reencountered. Mesanthura occidentalis Menzies and Barnard Mesanthura occidentalis Menzies and Barnard, 1959:15, fig. 9A, B.—Schultz, 1969:109, fig. 152a—d. 2 Mesanthura sp. of Brusca, 1973:202, fig. 7.17. The species (holotype 2 = 7 mm long) was collected from shallow water to water 55 m deep from Point Conception, California, to Bahia de San Quintin, Baja California. Brusca (1973) took specimens which he called “Mesanthura sp.” from algal mats in the intertidal zone of Puerto Penasco, at the northern part of the Gulf of California. Only limited mor- phological data were given, but some comparison of it was made with M. occidentalis. No formal name was given to the specimens however. Anthurid of Menzies Anthurid n. gen. & n. sp. Menzies, 1962:339. The 3 specimens on which this name is based were never formally de- scribed by Menzies (1962). They have chewing mouthparts and are in the family Anthuridae near Kupellonura Barnard (1925) according to Menzies. The specimens were briefly described without any definite dis- tinguishing characters by Menzies. Cortezura, new genus Diagnosis —Blind. Mouthparts modified for chewing. Frontal margin of cephalon with medial process as long as anterolateral corners. Peraeonal segment VII shortest of 7 peraeonal segments. Pleon with 6 conspicuous segments. Telson, not indurate, with uropods fan-shaped. Dorsum smooth, medial ventral ridge on all peraeonal segments. Antennae 1 and 2 short with VOLUME 90, NUMBER 4 843 few flagellar articles in both males and females. Maxillipedal palp of 3 arti- cles. Peraeopod I subchelate. Carpus underrides propodus only slightly on anterior peraeopods, never on posterior peraeopods. Pleopods 1 operculate. No statocysts. Etymology and gender.—The generic name is derived from the alternate name of the Gulf of California, “The Sea of Cortez.” The suffix “-ura” is without meaning. The gender is feminine. Type-species.—Cortezura penascoensis, sp. nov. Cortezura penascoensis, new species Description.—Blind. Cephalon slightly narrower and just over % length of peraeonal segment I; frontal margin with medial process as long as anterolateral corners. Peraeonal segments I-VI about equal in length; peraeonal segment VII shortest, about % length of peraeonal segment VI. Pleon with 6 conspicuous segments; telson slightly longer than 6 pleonal segments combined, with abruptly narrowing lateral borders ending in narrow truncate border. Dorsum smooth; no dorsal ornamentations, pits or lateral folds on any peraeonal segment or cephalon. Strong medial ridge ventrally; top of ridge flattened, with shallow medial groove along com- plete length. Antennae | and 2 short, similar in males and females. Antenna 1 with apparently only 2 flagellar articles; antenna 2 longer than antenna 1 with apparently only 1 flagellar article. Maxillipedal palp with 3 articles (4 if after Barnard, 1925:112); long setae on tiny apical article; at least 8 long setae on outer side of subapical segment; no endite apparent; exopod small, ovate, set into basal segment. Mandibles with tiny crenulations on incisor border; palp of 3 articles, apical article minute with one long seta; 6 long setae on article 2; 1 seta on basal article. Exopod of maxilla 1 with apex curved medially with many small teeth. Peraeopod I subchelate with large apical and smaller subapical claws. Large ovate propodus with inner margin slightly curved in female; with con- spicuous notch and process on male. Dactylus in male also with process on inner margin. Few setae on inner margin of segments of peraeopod | of male and female; many setae on inner margins of segments of peraeopod Il of male and female. Peraeopods II and III with carpus somewhat —underriding propodus. Carpus of each peraeopod from IV-VII slightly longer with carpus of VII longest. On peraeopods IV-VII carpus not underriding propodus. Pleopods 1 operculate; endopods only slightly narrower proximally than exopods. Long plumose fringing setae on apex of each ramus. Endopod of male pleopod 2 with laterally curved endopod (male stylet) on inner margin ending in rounded apex. Many long fringing setae on margins of other pleopods. Uropods with many long stiff fringing setae; apex of 844 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Figs. 1-8. Cortezura penascoensis, holotype male. 1, Anterior part; 2, Antenna 1; 3, Antenna 2; 4, Pleon and telson (dorsal view); 5, Pleon and telson (lateral view); 6, Peraeopod I with detail of palm; 7, Peraeopod II; 8, Peraeopod VII. VOLUME 90, NUMBER 4 845 Figs. 9-16. Cortezura penascoensis. 9, Pleopod 2 male; 10, Lateral view of female; 11, Peraeopod II female; 12, Peraeopod I female; 13, Maxillipedal palp; 14, 15, Mandibles; 16, Maxilla I. 846 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON exopod ovate; endopod longer than basis extending to about posteriou mar- gin of telson. No statocysts. Length to 10.5 mm. Peraeon VII is smaller and with undeveloped peraeopods in juveniles. Otherwise they are like adults. The smallest specimen (sex undetermined) with 7 mature peraeopods was 4.1 mm long. Type-locality and ecology.—The specimens were taken from a beach at Puertecitos, near Puerto Penasco, Sonora, Mexico, with amphipods on 13 April 1973. Males (3), females (15), unsexed (19) and juveniles (7) were collected (total 44). No females were gravid. Etymology—The name penascoensis means “from Pefiasco,’ the large town near which the specimens were caught. Remarks.—The species can be distinguished from all other anthurids from the west coast of North America except Bathura luna Schultz by the lack of eyes. It can be distinguished from B. luna in that B. luna has many flagellar articles on both antennae in contrast to the few on the flagella of the new species. Disposition of specimens.—The type-specimens have been deposited in the National Museum of Natural History (Smithsonian Institution) (holo- type 6 USNM 170410; allotype 2 USNM 170411; 25 paratypes 6 USNM 170412, 22 and juveniles USNM 170413). Other paratypes have been re- tained by the author. Discussion Bathura luna Schultz (1969) has been collected in deep water (783-812 m) and Silophasma geminatum (Menzies and Barnard, 1959) has been col- lected from shallow water to water 465 m deep. All other species includ- ing the new one described here are from less than 91 m, most of them be- ing collected from intertidal to relatively shallow water. The 8 species from the west coast are in 2 families. Silophasma geminatum (Menzies and Barnard) and Paranthura elegans Menzies are both in Paranthuridae of Menzies and Glynn (1968) (apex of maxillipedal palps pointed; modified for sucking). All others are in Anthuridae (apex of maxillipedal palps rounded; modified for chewing). Apparently the species with pointed maxillipedal palps are associated with plants (algae) and are adapted to feed on them. The rounded mouthparts are of more conventional detritus feeders. The species of the suborder Anthuridea were last classified by Barnard (1925). Many species from many localities in the world have been added since then and the group is badly in need of revision. When the revision is made, many of the genera included by Barnard will be restricted in def- inition and many new genera will be described. The revisions will un- doubtedly include more name changes among the species from the west coast of North America. VOLUME 90, NUMBER 4 847 Key to the Named Anthurids from the West Coast of North America la. With 6 large peraeonal segments Califanthura squamosissima lb. With 7 large peraeonal segments ys 2a. Blind 3 2b. With ocelli or eyes 4 3a. With many flagellar articles on antennae Bathura luna 3b. With 2 or less flagellar articles on antennae Cortezura penascoensis 4a. Pleon with conspicuous pleonal segments 5) 4b. Pleon entire 6 da. Apex of maxillipedal palp pointed Paranthura elegans Sb. Apex of maxillipedal palp rounded Apanthura californiensis 6a. Apex of maxillipedal palp pointed Silophasma geminatum 6b. Apex of maxillipedal palp rounded i 7a. Apex of maxillipedal palp with small rounded segment Cyathura munda 7b. Apex of maxillipedal palp pointed (but not modified for suck- ing) and produced medially Mesanthura occidentalis Acknowledgment The specimens of the new species were kindly supplied by Dr. Deborah M. Dexter, San Diego State University, and I thank her for the opportunity to describe them. Literature Cited Bamard, K. H. 1925. A revision of the family Anthuridae (Crustacea Isopoda). Jour. Linnean Soc. (Zool.) 36:109-160. Brusca, R. C. 1973. A handbook to the common intertidal invertebrates of the Gulf of California. Univ. Arizona Press, Tucson, Arizona, 427 pp. Iverson, E. W. 1974. Range extensions for some California marine isopod crustaceans. Bull. S. California Acad. Sci. 73(1):164—-169. Menzies, R. J. 1951. New marine isopods, chiefly from northern California, with notes on related forms. Proc. U.S. Nat. Mus. 101(3273): 105-156. 1962. The marine isopod fauna of Bahia de San Quintin, Baja California, Mexico. Pacific Nat. 3(11):337-348. , and J. L. Barnard. 1959. Marine Isopoda on coastal shelf bottoms of southern California: Systematics and Ecology. Pacific Nat. 1(11):1—43. , and P. W. Glynn. 1968. The common marine isopod Crustacea of Puerto Rico: A handbook for marine biologists. Stud. Fauna Curacao and Other Carib- bean Isl. 27:1-133. Poore, G. C. B. 1975. Australian species of Haliophasma (Crustacea: Isopoda: An- thuridae). Rec. Australian Mus. 29(19):503-533. Richardson, H. 1902. The marine and terrestrial isopods of the Bermudas, with descriptions of new genera and species. Trans. Connecticut Acad. Sci. Ll: 277-310. Schultz, G. A. 1964. Some marine isopod crustaceans from off the southern Cali- fornia coast. Pacific Sci. 18(3):307-314. 848 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON —. 1966. Marine isopods of the submarine canyons of the southern Californian continental shelf. Allan Hancock Pacific Exped. 27(4):1-56. —. 1969. How to know the marine isopod crustaceans. Pictured Key Nature Series. Wm. C. Brown Co. Pub., Dubuque, Iowa, viii + 359 pp. PROC. BIOL. SOC. WASH. 90(4), 1977, pp. 849-876 TWO NEW SPECIES OF ASHMUNELLA FROM DONA ANA COUNTY, NEW MEXICO, WITH NOTES ON THE ASHMUNELLA KOCHII CLAPP COMPLEX (GASTROPODA: PULMONATA: POLYGYRIDAE) Artie L. Metcalf and Richard A. Smartt Abstract.—Metcalf, A. L., Department of Biological Sciences, University of Texas at El] Paso 79968; and Smartt, R. A., National Fish and Wildlife Lab- oratory, Department of Biological Sciences, University of New Mexico, Albuquerque 87131.—Two new species of the polygyrid land snail genus Ashmunella are described from Dona Ana County, New Mexico: A. harrisi from the San Andres Mountains and A. todseni from the Organ Mountains. These species are judged to belong to the group of Ashmunella kochii Clapp, 1908. Additional species of this complex are enumerated and some aspects of their phylogeny and evolution are discussed. Introduction Two new species of the land snail genus Ashmunella from the San Andres and Organ Mountains, New Mexico, are described herein. Rela- tionships of the species of Ashmunella in these ranges and those of the Franklin Mountains, Texas, are discussed. The San Andres and Organ Mountains are located mainly in Dona Ana County, New Mexico, although the northern end of the San Andres range extends into Sierra County, New Mexico. Topographically, the two ranges are continuous, although, geographically, they are separated by San Augustin Pass (1,743 m), east of Organ, New Mexico (Fig. 1). The Franklin Mountains are directly south of the Organ Mountains in El Paso County, Texas. The San Andres Mountains are ca. 120 km long, with their highest point on Salinas Peak (2,730 m) in the northern end of the range (Fig. 1). The southern three-fourths of the range are comprised of completely faulted and tilted blocks, predominantly of Paleozoic limestones. Most mountains in this southern section are 2,100-2,280 m high, with San Andres Peak, the highest, reaching 2,510 m. Although the upper slopes of Salinas Peak are forested, montane forest is lacking in the southern part of the range. Scattered Pinyon Pine (Pinus edulis Engelm.), One-seeded Juniper (Junip- erus monosperma (Engelm.)), scrub oaks (Quercus spp.) and other small trees and shrubs occur in canyons and on higher slopes. Members of the Ashmunella kochii Clapp complex, discussed herein, inhabit the southern part of the range. 850 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON ——Thurgood Cn Sheep Mt. Salinas Pk Ladybug Pk —_ Rhodes Cn Ash Cn Sulphur Cn Kaylor Mt. Lost Man Cn Dead ManCn We Gardiner Pk - Mayberry Cn Ropessnr es San Andres Pk Salt Cn Black Brushy Mt om | Goat Mt ~ Bear Cn Black Mt San Augustin Pass 10 mi. 16.09 km VOLUME 90, NUMBER 4 851 Monzonitic igneous rocks predominate in the northern part of the Organ Mountains where impressive spires and pinnacles have formed. Prominent among these features are “Rabbit Ears,” “The Needles,” and “Organ Needle,” this latter peak being the highest in the range, at 2,727 m. The southern part of the range is predominantly of rhyolitic igneous rock with the highest peak being Organ Peak (2,704 m). The Organ Mountains support more forest than do the San Andres. Ponderosa Pine (Pinus ponderosa Laws.), Alligator Juniper (Juniperus deppeana Steud.) and Gambel Oak (Quercus gambelii Nutt.) are dominant species. The Franklin Mountains are a low, narrow, arid range, ca. 25 km long. Their highest peak, North Franklin Mountain, reaches 2,190 m in elevation. Mollusks of this range were discussed in Metcalf and Johnson (1971). In all the above mountains, accumulations of rock talus are the com- monest habitats of Ashmunellas. Limestone and rhyolitic formations are the major producers of such talus. Unless otherwise noted, terminology of shell characters follows that of Burch (1962: Figs. 11-13). Localities (abbreviated as “Loc.” or “Locs.”) noted in text are listed at the end of the paper. Criteria utilized in taxonomic allocation of species are conchological, conforming to those previously used in this genus. Where kinds with markedly different shells inter- grade, these have been treated as hybrids. These taxonomic assessments may eventually prove inadequate but seem to be useful at present in dis- cussing the fauna of a region poorly known malacologically. Abbreviations used in reference to types and paratypes are: ANSP = Academy of Natural Sciences of Philadelphia; DelIMNH = Delaware Mu- seum of Natural History; DMNH = Dallas Museum of Natural History; UA = University of Arizona; USNM = National Museum of Natural History, Smithsonian Institution, UTEP = University of Texas at El Paso. We are unaware of museum specimens of the new species described other than those listed below. Descriptions of New Species Ashmunella harrisi, new species Plate I, Figs. a, d; Fig. 2A, B Description of shell of holotype.—Shell thin, glossy, depressed, spire low, forming angle of 160°, sharply angular peripherally, with angularity < Fig. 1. General overview of San Andres and Organ Mountains, New Mexico, in- dicating major canyons, peaks and some other features. Lower contour line is drawn at 5,000 ft (1,524 m) and upper contour line is drawn at 7,000 ft (2,134 m). Thus, ele- vations in black are above 7,000 ft. x = localities at which Ashmunella pasonis pasonis was taken. 852 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON slightly above mid-height; body whorl not descending, umbilicus greatly ex- panding on body whorl; aperture oblique, at angle of 50° to vertical; upper, outer and lower lip thin, moderately flared, only slightly reflected, bearing a rounded palatal tooth compressed obliquely to lip, rising 0.7 mm above lip callus; palatal tooth separated by 1.4 mm from upper of 2 basal teeth, which are laterally compressed, 1.1 mm apart, both rising 0.6 mm above lip callus, the upper basal tooth slightly longer. Parietal callus thin, trans- parent (no thickening at edge), bearing an oblique tooth at angle of 40° to vertical, descending gradually anteriorly to form a vertical oriented “tail” and giving rise at inner upper end to a low, indistinct, horizontal branch 1.4 mm long (not white, as in oblique part of tooth). Embryonic half whorl smooth, minute growth wrinkles appearing near suture on remainder of embryonic whorl and gradually grading to low, smooth, regularly spaced growth wrinkles on remaining dorsal surface. Periphery of angularity slightly thickened but not keeled; ventral surface with low, widely spaced growth lines. Shell pale grayish-tan except for whitish lip and denticles. Variation in shells—Range of variation for some characters in 25 para- types from the type locality is indicated in Table 1. Shells vary con- siderably in size with a difference of 3.55 mm between diameters of largest and smallest mature shells. The upper branch of the parietal tooth is, at best, a low, indistinct swelling and in some specimens is absent. Shape and relative size of other denticles is uniform. A fulcrum is lacking in all speci- mens. Soft anatomy.—The following description derives from 5 paratypes that were dissected. General dorsal aspect of animals dark gray to blackish but with considerable variation among specimens. Two speci- mens blackish dorsally from tentacles to end of tail, grading to slightly lighter ventrally (sulci between raised areas lighter). One specimen with only tentacles and stripes extending posteriorly from them blackish, with remainder of body above sole grading from dark gray above to light gray below. Two other specimens intermediate between the two patterns of pigmentation described. Sole gray, lighter medially in some specimens. Mantle unpigmented except for some pale, minute, brown punctulations found (1) anteriorly alongside capillaries and (2) along and anterior to left afferent branchial blood vessel. Penis relatively long for genus with upper sac elongate, slender, narrower than lower sac, with 2 oblong patches of granular tissue, upper sac slightly wider than epiphallus (Fig. 2A, B). Vagina relatively long and slender, free oviduct %—% length of vagina. Spermatheca almost as long as uterus plus albumen gland. Measurements (all are lengths of organs and in mm) are as follows for specimens dissected (mean outside, range inside parentheses): Penis, 3.68 (3.14.8); Upper sac of penis, 2.27 (1.8-2.8); VOLUME 90, NUMBER 4 853 Table 1. Some measurements (in mm) for holotype and 25 paratypes (from type- locality) for each of 2 new species of Ashmunella. Length of parietal tooth refers to lower branch only. Palatal tooth is measured parallel to peristome. For each entry under paratypes, upper numerals indicate extremes and lower numerals mean and standard deviation (latter in parentheses). Ashmunella harrisi Ashmunella todseni Holotype Paratypes Holotype Paratypes Shell diameter 16.6 13.7-17.25 eS 11.6-14.3 15.69(.907 ) 12.80(.679) Shell height 5.7 4.45.9 5.4 Bll 5.27 (.379) 5.66(.421) Number of whorls 5.65 5.2—5.75 5.0 4.85-5.65 5.47(.164) 5.19(.196) Height of aperture 4.6 3.6—4.7 Boe 3.24.4 4.20(.289 ) 3.92(.281) Width of aperture 5.5 4.9-6.0 5.4 4.8-5.6 5.42.(.331) 5.14(.240) Width of umbilicus 4.4 3.4—4.9 Omi 20229 4.06(.374) 2.42 (.205) _ Parietal tooth length 2.8 23=3.1 2.5 2.4—3.0 2.62 (.204) 2.67 (.149) Palatal tooth length 1.0 0.8-1.9 1E2 OS ea | 1.24(.274) 1.30(.204) _ Width of reflected part of lip 1.0 0.8-1.5 1.6 0.9-1.4 | 1.22(.191) 1.16(.138) _ Epiphallus, 27.45 (24.6-31.5); Vas deferens, 29.37 (22.0-33.1); Spermatheca plus duct, 30.77 (26.4-35.4); Vagina, 4.85 (4.4-5.7); Free oviduct, 2.05 (1.8-2.5). Diagnosis —An Ashmunella with genitalia typical of the A. kochii group (discussed below): differentiable from other members of the group by the combination of thin, glossy, smooth shell, absence of fulcrum and with no or an extremely reduced upper branch of the parietal tooth. It is far smaller than A. k. kochii, its neighbor to the northwest. Populations judged _ to be hybrids between A. harrisi and A. pasonis pasonis occur in the eastern part of Bear Canyon. These are discussed below. Etymology.—The species is named in honor of Dr. Arthur H. Harris, University of Texas at El Paso, who first collected the species. Distribution —Ashmunella harrisi is known from 2 canyons on_ the east side of Goat Mountain in the southeastern part of the San Andres 854 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Mountains (Locs. 10 and 11). The type-locality is Loc. 10, with the holotype and associated paratypes collected on 11 April 1975. Disposition of types——Holotype, ANSP 340724. Paratypes: ANSP 340725; DeIMNH 106682; DMNH 4535; UA 6218; USNM 758527; UTEP 3139, 4413. Ashmunella todseni, new species Plate) Figs) baie. hice JC Das Description of shell of holotype.—Shell thick, depressed, spire low, form- ing angle of ca. 150°, sharply angular (subcarinate) peripherally, with an- gulation approximately midway of height. Five whorls, body whorl not descending; umbilicus moderately narrow, 3.78 times in shell diameter. Aperture markedly oblique, at angle of 50° to vertical. Upper, outer and lower margin of peristome rounded, lip broad and reflected, outer re- flected part of lip thin, inner part thickened, calluslike, especially at upper end, where a thick callus extends back 0.7 mm along roof of body whorl. Lip continuous with thickened, anterior, raised edge of parietal callus, raised edge of callus white, remainder of callus translucent but bearing large oblique parietal tooth; lower % of tooth parallel to raised anterior parietal callus margin, upper 7% of tooth at angle of 115° to lower part, slightly sinuous; upper end of parietal tooth giving rise to short, low, arcuate branch. Lip peristome giving rise to rounded palatal tooth, ris- ing 0.5 mm above callus and 2 compressed basal teeth, the uppermost slightly larger and rising 0.65 mm, the lower only 0.5 mm above callus; all 3 lip teeth evenly spaced, ca. 1 mm, from each other. Embryonic half whorl slightly roughened, surface of whorls 0.5-1.75 finely wrinkled, re- mainder of dorsal surface with regularly spaced, moderately strong growth wrinkles. Peripheral angularity slightly thickened, almost keeled. Ventral surface with low, regularly spaced growth wrinkles, becoming stronger near aperture. Rows of minute cuticular “hairs” on walls of umbilicus. Shell light reddish brown except for strongly contrasting whitish lip, teeth and raised margin of parietal callus. Variation in shells—Range of variation in shell characters for 25 para- types from the type locality is indicated in Table 1. The anterior edge of the parietal peristome varies from appressed to strongly raised. The oblique parietal tooth varies in length and height, being relatively massive in some specimens. The upper branch of the parietal tooth is, at best, a weak swelling and is absent in some specimens. The palatal tooth varies from short and rounded to longer and rectangular. A fulcrum is lacking in all specimens. Cuticular hairs and scales occur on the entire ventral surface and on the dorsal surface of younger whorls in specimens from Loc. 21. Soft anatomy.—(Fig. 2C, D, E). The following description is derived from 5 paratypes dissected. Head, including all tentacles, uniformly black. Posteriorly and ventrally on body, color lightening but with raised areas Ol VOLUME 90, NUMBER 4 85! B Fig. 2. A and B, Genitalia of Ashmunella harrisi,; C, Dorsal view of mantle of Ashmunella todseni; D and E, Genitalia of A. todseni. ag = albumen gland; e = epiphallus; f = flagellum; fo = free oviduct; fp = fertilization pouch or talon; go genital orifice; Is = lower sac of penis; od = ovotestis or hermaphroditic duct; og oviducal gland; pg = prostate gland; pr = penial retractor muscle; s spermatheca; sd = spermathecal duct; us = upper sac of penis; v = vagina; vd = vas deferens. 856 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON darker than intervening sulci. Margin and sole of foot uniformly light gray. Collar of mantle gray with dark brown margin. Mantle especially well pigmented for members of genus (Fig. 2C) with area immediately pos- terior to collar prominently flecked and streaked with black pigment. Ef- ferent and left afferent branchial blood vessels and their capillaries out- lined with black punctulations. Kidney outlined with interrupted dark punctulations. Mantle pigmented dorsally on part or all of first 2 whorls and atop third whorl with 2 rows of dark punctulations with ladderlike cross-connections. Penis with upper sac narrower than lower sac; upper sac with 2 oblong patches of granular tissue visible externally (Fig. 2E). Free oviduct ca. one-half length of vagina. Spermatheca extending beyond uterus and al- bumen gland. Fertilization pouch (= talon) comprising 2 or 3 globular bodies. Genitalia are shown in Fig. 2D, E. Measurements of genital organs for the specimens dissected (given as in A. harrisi, above) are: Penis, 3.36 (3.2-3.5); Epiphallus, 24.8 (22.6-28.2):; Vas deferens, 24.9 (21.5-26.1); Spermatheca plus duct, 31.48 (27.2-35.2); Vagina, 5.1 (4.9-5.3). Diagnosis —An Ashmunella with genitalia typical of the A. kochii group but with mantle more heavily pigmented than in other species of the group. Shells differentiable in being subcarinate with large, sinuous lower and rudimentary or no upper parietal denticle, and lacking fulcrum. Its nearest neighbor, A. organensis Pilsbry, 1936, has greatly reduced dentic- ulation and rounded, rather than subcarinate, shells. Ashmunella auriculata Vagvolgyi, 1974, also of the Organ Mountains, has a thinner, smoother, angular shell with a wider, shallower umbilicus. Etymology.—This species is named in honor of Dr. Thomas K. Todsen, White Sands Missile Range, New Mexico, to whom we are especially in- debted for escort provided on the Missile Range, for his knowledge of the area and for his good companionship. Distribution —Ashmunella todseni has been collected only in Maple and Texas Canyons in the northeastern part of the Organ Mountains (Locs. 21, 22, 24). The type-locality is in Maple Canyon, Loc. 24, with holotype and associated paratypes collected on 24 May 1975. At this time it is not possible to ascertain to what extent A. todseni may extend around the southeastern part of the Organ Mountains from the localities listed here. This area, mainly on Fort Bliss Military Reservation, is used as an artillery range and access is not authorized. Habitat—cologically, A. todseni may differ from the other species of Ashmunella in the Organ Mountains. At its type-locality, where especially abundant, it was found only in the upper 10 cm of small, rubbly talus of igneous rock. At lower depths, where other species of Ashmunella in the Organ Mountains are usually found, A. todseni did not occur. It seems VOLUME 90, NUMBER 4 857 Fig. 3. Distal region of genitalia of some species of Ashmunella from southern New Mexico and adjacent Texas. Abbreviations as in Fig. 2. A, A. organensis (Loc. 29); B, A. amblya amblya (Pine Spring Canyon, Guadalupe Mts., Texas); C, A. amblya cornudasensis (Flattop Mt. of Cormnudas Mts., New Mexico); D, A. kochii kochii (Loe. 7); E, A. kochii sanandresensis (Loc. 6); F, A. pasonis pasonis (Loc. 1); G, A. pasonis pasonis (Loc. 36); H, A. auriculata (Loc. 18). 858 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON || to be, then, a species especially well adapted to an arid microhabitat, prob- ably subject to pronounced extremes of temperature. Disposition of types—Holotype, ANSP 340726. Paratypes: ANSP | 340727; DeIMNH 106683; DMNH 4534; UA 6217; USNM 758528; UTEP 4476, 4744, 4746. Additional Species of the Ashmunella kochii Complex Including the 2 species described above, 7 species of Ashmunella are known from the San Andres, Organ and Franklin Mountains. Ashmunella salinasensis Vagvolgyi, 1974, occurs on Salinas Peak in the northern part of the San Andres Mountains. It clearly belongs to the group of Ashmunella rhyssa (Dall, 1897), a complex widespread in the Sacramento and Sierra Blanca Mountains on the east side of the Tularosa Basin (Metcalf, 1973:38). Similarities in morphology of genitalia suggest that the remaining 6 spe- cies of the mountains treated here belong to a single complex, termed the “Ashmunella kochii complex,” in reference to the first-named member of the group (Clapp, 1908). In this group the penis is of the kind described by Pilsbry (1940:913, 914): “the upper part of the penis is only about half as wide as the lower, and sometimes hardly distinguishable from the epiphallus.” (See Figs. 2; 3A, D-H). Also belonging to the A. kochii complex are (1) A. rileyensis Metcalf and Hurley, 1971, from Mount Riley, 40 km west of the Franklin Mountains (in New Mexico) and (2) A. kochii caballoensis Vagvolgyi, 1974, from the Caballo Mountains, ca. 55 km west of the San Andres Mountains. Remaining taxa of the A. kochii group in the San Andres, Organ and Franklin Mountains are treated below. Some shell characters utilized in delimiting taxa of Ashmunella in these mountains are given in the ac- companying key. (Upper parietal denticle = upper branch of parietal den- ticle.) A Key to Shells of the Taxa of the Genus Ashmunella of the San Andres, Organ and Franklin Mountains la. Shell heavily ribbed A. salinasensis Vagvolgyi lb. Shell not heavily ribbed 2 2a. Parietal denticles fused into a subtriangular or subquadrate placque that occupies most of parietal wall 3 2b. Parietal denticles not forming placque on parietal wall 4 3a. Shell keeled, bearing cuticular scales A. pasonis pasonis (Drake) 3b. Shell angular rather than keeled; lacking cuticular scales A. pasonis polygyroidea Vagvolgyi 4a. Fulcrum (internal denticle on floor of body whorl) lacking ) 4b. Fulcrum present 8 VOLUME 90, NUMBER 4 859 5a. Body whorl subcarinate peripherally; adult shells bearing cuticular hairs in umbilical region A. todseni n. sp. Sb. Body whorl angular or rounded peripherally; adults with no cuticu- lar hairs 6 6a. Parietal and lip denticles rudimentary or absent A. organensis Pilsbry 6b. Well developed denticles present both in lip and on parietal wall i 7a. Upper parietal denticle well developed A. auriculata Vagvolgyi 7b. Upper parietal denticle absent or rudimentary A. harrisi n. sp. Sa. Shells larger than 18.5 mm in diameter; upper parietal denticle rudimentary or absent A. kochii kochii Clapp Sb. Shells smaller than 18.5 mm in diameter; upper parietal denticle well developed A. kochii sanandresensis Vagvolgyi Ashmunella kochii kochii Clapp ews Ibe lie, dD, ILoes, Wate!) Ashmunella kochii G. H. Clapp, 1908:77; Pl. 8, figs. 1-3. Clapp (1908) described the type-locality of A. kochii thus: “Black Mountain, at the southern end of the San Andreas Range, Donna Ana county, New Mexico, at an elevation of about 6,800 feet.” Pilsbry (1940:977) named a subspecies, A. kochii amblya, from the Guadalupe Mountains of New Mexico and Texas, ca. 180 km to the east-southeast. Vagvolgyi (1974: 143) suggested that Clapp’s type of A. k. kochii might actually have come from the Guadalupe rather than from the San Andres Mountains. The problem of type-localities is further complicated by the existence of 2 mountains indicated as “Black Mountain” on maps of the southern San Andres Mountains. The more southern of these is located south of Bear Canyon (Figs. 1, 6) and centers around the northeast corner of Sec. 1, T. 21S, R. 4E. The more northern peak is located on San Andres National Wildlife Refuge, south of Salt Canyon (Figs. 1, 6) and centers around Sec. 3, T. 20S, R. 4E. Mr. John Kiger, manager of the Refuge (pers. comm.) has informed us that the more northern mountain is locally called “Black Brushy Mountain” and we shall use that name here. On the southern and northern sides of (the southern) Black Mountain, we have collected, respectively, A. pasonis pasonis (Loc. 16) and hybrids between A. p. pasonis and A. harrisi (Locs. 13-15). At Loc. 7, on the northern side of Black Brushy Mountain, we have collected a large Ashmunella, shells of which seem indistinguishable from the type of A. k. kochii Clapp. We think it most likely, then, that the type of A. k. kochii did come from the southern San Andres Mountains, as originally indicated by Clapp, and probably from Black Brushy Mountain. It is understandable that Vag- volgyi (1974:143) considered the Guadalupe Mountains to be the type- 860 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Plate I. Shells of some kinds of Ashmunella from southern New Mexico and adjacent Texas. Measurements are for diameter of shell. a—c, viewed from side, toward aperture; d-1, umbilical view. a, d, Holotype of A. harrisi, ANSP 340724 (16.6 mm); b, e, Holotype of A. todseni, ANSP 340726 (13.5 mm); c-1 and f-1, A. pasonis pasonis, Loc. 1 (12.45 mm); c-2 and f-2, A. pasonis pasonis, Loc. 36 (12.45 mm); c-3, A. pasonis polygyroidea, Loc. 42 (12.9 mm); g and h, hybrids of A. pasonis pasonis x A. harrisi from Loc. 15 (g—16.2 mm) and Loc. 14 (h—14.5 mm); i, A. kochii sanandresensis from VOLUME 90, NUMBER 4 861 locality because the shells of A. k. kochii from Black Brushy Mountain (not seen by him) and those of A. k. amblya from its type-locality in Pine Spring Canyon in the Guadalupe Mountains are strikingly similar (PI. I, j, k). There are slight differences, however. The (lower) parietal tooth of A. k. amblya is straighter and more rectangular in shape (as shown in Pilsbry, 1940: Fig. 566d), whereas in specimens from Black Brushy Mountain this tooth is slightly sinuous and incurved basally to form a “tail” (op. cit., Fig. 566a, b). On the average, the palatal tooth of shells of A. k. amblya is shorter and more rounded than in specimens from Black Brushy Mountain. The genitalia of the 2 kinds show marked differences. The upper sac of the penis is unusually short, for the genus, and much broader than the epiphallus in A. k. amblya (Fig. 3B). In specimens from Black Brushy Mountain (Fig. 3D) the upper sac is long and only slightly broader than the epiphallus, as in other species from the southern San Andres and Organ Mountains. In view of the differences enumerated above, we propose that A. amblya be accorded specific rank. Vagvolgyi (1974) described A. kochii cornudasensis from the Cornudas Mountains, 70 km west of the Guadalupe Mountains and 115 km south- east of Black Brushy Mountain. This Ashmunella has genitalia (Fig. 3C) of the kind seen in A. amblya. Thus, we propose that 2 subspecies be recog- nized in A. amblya: (1) the nominal subspecies from the Guadalupe Mountains and (2) A. ambyla cornudasensis Vagvolgyi from the Cornudas Mountains. Herein, we deem A. kochii to comprise 3 subspecies: (1) the nominal race (Black Brushy and Goat Mountains), (2) A. k. sanandresensis Vag- volgyi, 1974, from San Andres Peak, immediately north of Black Brushy Mountain, and (3) A. k. caballoensis Vagvolgyi, 1974, from the Caballo Mountains, west of the San Andres Mountains. Ashmunella kochii sanandresensis Vagvolgyi Plate I, i; Fig. 3E. Loc. 6 Ashmunella kochi amblya H. A. Pilsbry, 1940:977 (provisionally referred). Ashmunella kochi G. H. Clapp, 1908.—Metcalf, A. L. and P. A. Hurley, LOA 22. Ashmunella kochi sanandresensis J. Vagvolgyi, 1974:145; Pl. I, 3, 3a; fig. Ac. Shells of this race from near its type-locality on the western slope of San Andres Peak differ from A. k. kochii in being smaller and having a < Loc. 6 (17.1 mm); j, A. amblya amblya from Pine Spring Canyon, Guadalupe Moun- tains, Texas (20.9 mm); k, A. kochii kochii from Loc. 7 (19.6 mm); 1, Hybrid of A. p. pasonis & A. kochii sanandresensis from Loc. 4 (13.95 mm). 862 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON well developed upper parietal tooth (see Pl. I, i, k). No living intergrades between the 2 subspecies were found in the lower mountains between San Andres Peak and Black Brushy Mountain. However, on the south wall of — Ash Canyon (32°37’50’N; 106°32’18’W) occur fossils (probably late Pleis- tocene) that seem to be intergrades. These are of the smaller size of A. k. sanandresensis but lack an upper parietal tooth. In the northwesternmost — part of San Andres Peak (Locs. 4, 5), A. k. sanandresensis shows evidence of hybridizing with A. pasonis pasonis, which occurs in the mountains to the north. Ashmunella pasonis pasonis (Drake) | Pl. I, c-1, c-2, f-1, f£-2; Fig. 3F, G; Fig. 4A, B. Locs. 1-3, 9, 16, 34-40 Polygyra pasonis R. J. Drake, 1951:44-46; Figs. 1-4. Ashmunella pasonis.—R. J. Drake, 1952, Ibid., 9:30. Drake (1951) described Polygyra pasonis from shells collected in Vinton Canyon in the northern Franklin Mountains, E] Paso County, Texas. He later (1952) reassigned P. pasonis to the genus Ashmunella. Vagvolgyi (1974:156) partitioned A. pasonis of the Franklin Mountains into a more northern, nominal subspecies and a more southern subspecies that he named A. pasonis polygyroidea. Pleistocene fossils, closer in shell morphology to A. p. polygyroidea than to the nominal subspecies, have been found in the northern part of the Franklin Mountains where A. p. pasonis presently occurs. This indicates (1) that in an earlier Quaternary time probably all of the range was inhabited by A. p. polygyroidea and (2) in subsequent time, populations in the northern part of the range have been evolving toward the shell morphology of A. p. pasonis. Shells of living populations demonstrate intergradation from north to south. Salient characters show- ing such clinal intergradation include (listing the more plesiomorphic con- dition of A. p. polygyroidea first and the more apomorphic A. p. pasonis second): (1) shell higher and more rounded peripherally to shell more de- pressed and carinate; (2) shell without to shell with cuticular scales; (3) smaller to larger lip teeth; (4) parietal placque that is smaller, more triangular and less raised anteriorly to placque that is larger, more quad- rate and more raised anteriorly; (5) lip less reflected and with less develop- ment of a “neck” behind the aperture to lip greatly reflected and with well developed “neck.” Apertural features distinguishing the 2 subspecies are shown in Fig. 4. The nominal subspecies of A. pasonis also occurs in the San Andres Mountains. It has been found in 2 areas there, one to the north and one to the south (see “x's” in Fig. 1). In the more northern area, A. pasonis pasonts occurs at least from Lost Man and Dead Man Canyons (Locs. 1, 2) south to Mayberry Canyon (Loc. 3) and it hybridizes with A. k. sanandresen- VOLUME 90, NUMBER 4 863 Fig. 4. Shells of Ashmunella pasonis from the Franklin Mts., El Paso County, Texas. A-D, Apertural areas showing graduation from apomorphic A. pasonis pasonis (A and B—Locs. 34 and 36) to plesiomorphic A. pasonis polygyroidea (C and D—Locs. 41 and 42); E, Shell with part of body whorl removed to show fulcrum (t). sis in the northern part of San Andres Peak (Locs. 4, 5). In the south, A. p. pasonis occurs in upper Little San Nicholas Canyon together with A. &. kochii (Loc. 9) with which it does not hybridize. The 2 species seem to oc- cupy slightly different habitats. In the southern area it also occurs on 864 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON a oledad Canyon VOLUME 90, NUMBER 4 865 Black Mountain (Locs. 12, 16), although on its northern slope in Bear Canyon (Locs. 13-15) it hybridizes with A. harrisi. Ashmunella pasonis polygyroidea Vagvolgyi Plate J, c-3; Fig. 4C, D. Locs. 41-42 Ashmunella pasonis R. J. Drake, 1951.—Metcalf, A. L. and W. E. Johnson, 1971:94, 100. Ashmunella pasonis polygyroidea J. Vagvolgyi, 1974:156, Pl. I, fig. 10. This race of A. pasonis presently is found living only in the central part of the Franklin Mountains. Pleistocene fossils indicate that it pre- viously inhabited the southern and northern parts of the range as well. Subsequently, it has become extinct in the southern part of the range and has seemingly evolved into A. pasonis pasonis in the northern part. Vag- volgyi (1974:157) suggested that A. p. polygyroidea might represent hybrids between A. pasonis pasonis and some race of A. kochii. However, the na- ture of the fossils of A. pasonis in the Franklin Mountains seems clearly to demonstrate that A. p. polygyroidea is the ancestor of A. p. pasonis and that its resemblance to A. kochii results from plesiomorphic features derived from an A. kochii-like ancestor. Ashmunella auriculata Vagvolgyi Fig. 3H. Locs. 17-20, 25, 26 Ashmunella mearnsii (Dall, 1895).—Cockerell, T. D. A., 1897:69. Ashmunella kochi G. H. Clapp, 1908.—Pilsbry, H. A., 1915:329.—Metcalf, A. L., 1969: Table 1.—Metcalf, A. L. and P. A. Hurley, 1971:122, 126. Ashmunella auriculata J. Vagvolgyi, 1974:150; Pl. I, figs. 5, 5a. Vagvolgyi (1974:150) indicated the type-locality of A. auriculata to be in Boulder Canyon in the southern part of the Organ Mountains. This species also occurs in the northern, monzonitic “Needles” area of the moun- tains, on Baylor Peak, and in the lower part of Fillmore Canyon (Fig. 5). Between these northern populations and the type-locality Ashmunella organensis occurs. < Fig. 5. Organ Mountains, Dona Ana County, New Mexico. Contours are drawn at 6,000 ft (1,829 m), 7,000 ft (2,134 m) and 7,500 ft (2,286 m). Thus, elevations in black are above 7,500 ft. Open circles = localities of Ashmunella todseni; filled circles = localities of Ashmunella organensis; X = localities at which Ashmunella auriculata was taken in this survey (type-locality is in Boulder Canyon). B Baylor Peak; N Organ Needle; P = Organ Peak; RE = Rabbit Ears. 866 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Ashmunella organensis Pilsbry Fig. 3A. Locs. 23, 27-33 Ashmunella organensis H. A. Pilsbry, 1936:101. Pilsbry (1936:101) described this species from an upper branch of Ice Canyon in the west-central part of the Organ Mountains. We have found it to occur in the central and southwestern parts of the range (Fig. 5) and even on the west wall of Boulder Canyon (Loc. 32), the canyon that Vagvolgyi (1974:151) recorded as the type-locality of A. auriculata. Ash- munella organensis is not known to hybridize with other species of the Organ Mountains (A. todseni and A. auriculata). In regard to shell char- acters it bears little resemblance to the A. kochii complex, having a high spire (more pronounced at higher elevations) and with rudimentary or no apertural teeth. However, morphology of the genitalia indicates al- liance with the group of A. kochii. Hybridization In 2 areas in the San Andres Mountains hybridization seems to occur between Ashmunella pasonis pasonis and another species, suggesting some genetic plasticity on the part of A. p. pasonis. 1. Ashmunella p. pasonis < A. harrisi. Pl. I, g, h; Locs. 13-15. Having a shell morphology judged to be among the most plesiomorphic of the A. kochii complex, it is surprising that A. harrisi hybridizes with A. p. pasonis, deemed the most apomorphic member of the complex in regard to shell morphology. A cline grading from one species to the other is shown in 4 collections taken along the south wall of Bear Canyon. which lies south of Goat Mountain (with type-locality of A. harrisi on its east slope) and north of Black Mountain (Fig. 6). At the easternmost locality (Loc. 15, Fig. 6) shells resemble types of A. harrisi except for the parietal area, in which the interdental callus has become thickened and an upper branch of the parietal tooth has become a salient feature (Pl. I, g). At Loc. 14, 2.4 km west-northwest of Loc. 15, the parietal callus has become greatly thickened and placquelike and the 2 branches of the tooth with the an- terior margin of the callus, form a triangular, raised rim around the placque. The outer lip has become slightly recurved with the result that lip teeth are more deeply set. A weak keel has appeared and cuticular scales cover much of the shell (PI. I, h). Shells from Loc. 13 (Fig. 6), 0.4 km northeast of Loc. 14 are still smaller, scalier and more flattened and keeled than those from Loc. 14. An anteriorly elevated parietal placque covers almost all the parietal wall. The lip is flared and recurved. These shells approach the shell type of A. p. pasonis and shells from Loc. 12, 0.39 km north of Loc. 13, seem indistinguishable from those of A. p. pasonis except tor lacking a fulcrum. Specimens of A. p. pasonis from Locs. 9 and 16 also lack the fulcrum (see Fig. 4E), which is found in this species in the Frank- VOLUME 90, NUMBER 4 867 °° ole S DONE | ° ‘ o°n [0 0,0 aca °° On Fig. 6. Environs of Goat Mountain and Bear Canyon, San Andres Mts., Dona Ana County, New Mexico. Numbers in italics are localities of collections; non-italicized numbers are of surveyed sections. H = type locality of Ashmunella harrisi; h another locality of A. harrisi; p = locality of Ashmunella pasonis pasonis; hp localities with intergrades between A. p. pasonis and A. harrisi. Contour lines are at 500 ft (152.4 m) intervals with black, dotted area above 7,000 ft (2,134 m) and open-dotted area 4,500—5,000 ft (1,372-1,524 m). lin Mountains and in northern populations of the San Andres Mountains (Locs. 1-3). Perhaps this lack of a fulcrum in southern populations of the San Andres Mountains reflects a genetic contribution from A. harrisi, which also lacks this structure. 868 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 2. Ashmunella p. pasonis < A. kochii sanandresensis. Pl. I, 1; Locs. 4-5. Typical specimens of A. p. pasonis occur in Lost Man, Dead Man and Mayberry Canyons in the San Andres Mountains (Fig. 1). The type- locality of A. k. sanandresensis is to the south in the central, western part of San Andres Peak near Loc. 6. In the northern part of San Andres Peak hybridization occurs between the two kinds (Locs. 4 and 5). Specif- ically, towards the north on San Andres Peak, there is thickening of the parietal callus between the upper and lower branches of the parietal tooth to produce a parietal placque approaching that, which is typical of A. p. pasonis. There is also an increase in relative size of the teeth of the outer lip. Shells become smaller towards the north, approaching the size of Aen pasonis (eleaiei): Phylogenetic Trends in Shells in the Ashmunella kochii Complex Genitalia of the members of the A. kochii complex exhibit much similarity. Shells, however, show considerable variability and may offer clues to phy- logeny within the group. Pilsbry (1905:255) hypothesized that, as regards denticulation in Ashmunella, a tridentate condition (teeth in palatal and basal positions on the lip and a tooth on the parietal callus) might be re- garded as primitive. From this presumed plesiomorphic tridentate con- dition some phyletic lines have evolved towards a loss of some or even all denticles while others have increased their armature. This increase seems generally to have taken place in 3 different ways: 1. In some lines the basal lip tooth has bifurcated to produce 2 teeth, subtended by a common callus; these teeth have, then, in subsequent evolution, become more widely separated. Ashmunella ruidosana Metcalf, 1973, a fossil species, and A. thomsoniana (Ancey, 1887) and A. pseudo- donta (Dall, 1897), both living species, seem to demonstrate a condition in which this bifurcation is taking place. 2. The parietal tooth seems, independently, in several lines, to have gained an upper branch oriented at an acute angle to the lower branch, thus yielding a V-shaped appearance. In a few species there has been, subsequently, a trend in which the parietal callus between the upper and lower arms of the tooth has become thickened. In the extreme ex- ample of A. pasonis the interdental callus has thickened greatly, becoming elevated anteriorly from the underlying shell wall and forming, in an earlier stage of evolution, a triangular placque as seen in A. p. polygyroidea (Fig. 4C, D). In subsequent evolution, producing A. p. pasonis, the placque has become even larger and more quadrate in shape (Fig. 4A, B). 3. In some species a tooth has appeared several mm behind the aperture on the inner basal part of the body whorl. This tooth has been termed a fulcrum by Vagvolgyi (1974: Fig. 1). A fulcrum is present in all spe- cies of the A. kochii complex except those inhabiting the Organ Mountains, in A. rileyensis, A. harrisi and in hybrids of A. pasonis x A. harrisi. VOLUME 90, NUMBER 4 869 Primitively, shells of Ashmunella were probably generally rounded peripherally and moderately high spired as in the shells of A. organensis from higher elevations. Evolution towards a more depressed and often carinate shell seems, somehow, to be related to living at lower (hence, more arid) elevations. Even specimens of A. organensis from lower ele- vations exhibit more depressed shells than their relatives at higher eleva- tions. Carination seems especially common in Ashmunella inhabiting talus of flat limestone rocks. Cuticular scales have developed in several lines of Ashmunella. Again, this seems to be most often exhibited by species living in lower, more arid habitats, as in A. lepiderma Pilsbry and Ferriss, 1910, in A. bequaerti Clench and Miller, 1966, and in A. todseni and A. p. pasonis. A Suggested History of the A. kochii Complex On the basis of present and past known distributions and on shell-mor- phological evidence provided by living and fossil materials, the follow- ing model is suggested. We present it to be tested against further dis- coveries and more refined analytical methods. 1. With its relatively high-spired shell and weak denticulation, A. or- ganensis seems to be somewhat removed from other members of the com- plex. It may long have inhabited the area. Fossils of A. hawleyi Metcalf, 1973, from deposits of probable mid-Quaternary age along the Rio Grande Valley resemble A. organensis as does also a single fossil taken from de- posits along Salt Canyon on the north side of Black Brushy Mountain in the San Andres range, northwest of Loc. 7. In younger deposits on the same hillslope occur fossils of A. kochii kochii, which presently inhabits this slope. There is, then, some evidence that A. organensis is an ancient and formerly more widespread member of the A. kochii complex, now restricted to the Organ Mountains. 2. The remaining species of the complex have dispersed widely in the mountain ranges treated here. A vicariance model could be employed to account for their initial occupation of the ranges; however, some instances of later dispersal by propagules seem likely, as suggested below. Prob- ably a plesiomorphic shell of this latter group would have 4 denticles, 3 in the outer lip and a parietal tooth (bearing, at most, a very weak upper branch), and lacked a fulcrum. Only A. harrisi and A. todseni retain this kind of denticulation today. These species may stem from a relatively early stage in the evolution of the A. kochii complex. Restricted in small eastern enclaves of their respective mountain ranges, one (harrisi) has re- tained a more conservative shell; the other (todseni) has evolved into a xeric-tolerant species with a more specialized shell. 3. A slightly more advanced shell might have acquired a stronger upper parietal branch but still lacked a fulcrum. Such a shell is possessed by A. auriculata and A. rileyensis. These species are similar and A. rileyensis 870 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON may have been derived from a propagule of A. auriculata from the Organ Mountains. 10622822 W .-SWa4,NHA,SW%4, Sec. 31, T: 20'S, R. 5 E. Talus of igneous rock on north-facing wall of Bear Cn. (N side of Black Mts) el Om. tomo 2200 Ne 106°28’52°W. S24. NEY NE, Sec. 12; T. 21S, R. 4 E. Lime- stone talus on walls of south-draining canyon on S flank of Black Mt. 1,828 m. Organ Mountains, Dona Ana County, New Mexico: 17. 32°23’41’N; 106°34’50”W. .08 km W of center, E section line, Sec. 13, T. 22 S, R. 3 E. From prominent rhyolitic talus accumulation on W side of Baylor Peak. 2,134 m. ISao2 22047N: 106°34730°W. SWANEYANW%4, Sec. 30, T. 22 S, RB. 4E. Monzonitic talus on NE slope of Rabbit Ears Peaks and directly below valley between the two “Ears.” 2,073 m. — 19. 32°21’29”"N; 106°34’14’”W. Area immediately S of center, N section line, Sec. 31, T. 22S, R. 4 E. In rock-soil mixture in grove of Gambel Oak on E side of “The Needles.” 2,225 m. 20a 20 2GN- 106%33/27 7 W. SW, NESW, Sec) 32, TT. 22S: RK. 4 E. Upper end of Indian Hollow Cn., below (east of) Organ Needle in talus-soil mixture in grove of Bigtooth Maple. 2,225 m. Both north and south of Baylor Peak and along the east side of The Needles, A. auriculata seems generally to occur where there is talus above 2,000 m in light forest of Gambel Oak and Ponderosa Pine. 21. 32°20'58’N; 106°30’59"W. SW%,NE%4,SE%, Sec. 34, T. 22S, R. 4 E. Talus accumulation near mines at upper, SW end of Texas Cn. 1,814 m. 22. 32°20'43’N; 106°30’41”W. SW%4,SW%4,SW*%, Sec. 35, T. 22 S, R. 4 E. Upper end of a S branch of Texas Cn., in “bowl” distinguished by tall Ponderosa Pines, under high cliffs, in talus. 1,860 m. 23. 32°20'39’N; 106°31’46”W. Northwesternmost corner of Sec. 3, T. 23 S, R. 4 E. Rhyolitic talus on west-facing wall of Rock Springs Cn. 1,920 m. 24, 32°20/26”"N; 106°31’2”W. NW%,SE%,NE%, Sec. 3, T. 23 S, R. 4 E. Talus accumulation of igneous rock on E side of E branch of Maple Cn. Type-locality of A. todseni. 2,042 m. 25. 32°20/11”N; 106°33’36’W. NE%,NW%4,SW%, Sec. 5, T. 23 S, R. 4 E. Rhyolitic talus on west-facing wall of “The Narrows” in Fillmore Cn. 2,164 m. 26. 32°20’9”N; 106°34’23”W. NE%,NE%,SW%, Sec. 6, T. 23 S, R. 4 E. Rhyolitic talus on north-facing wall of Fillmore Cn., 0.4 km SW of Fill- more Spring, below small cave, higher on canyon wall. 1,996 m. 27. 32°20'4’"N; 106°32/54”"W. SEY,NE%,SE%, Sec. 5, T. 23 S, R. 4 E. 874 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON | In leaf litter and loose talus on east-facing wall of narrow canyon, .15 km SE of Rock House Spring. 2,408 m. 98. 32°19'53’N; 106°32’56”“W. SW%4,SE*%{SE™%, Sec. 5, Ta 2ensner san | North slope of Organ Peak. 2,620 m. 99. 32°19'30’N; 106°34’31”W. NW%4,SE%,NW%, Sec. 7, T. 23 S, R. 4 E. Rhyolitic talus on south-facing wall of mouth of Ice Cn., .48 km NW of Dripping Spring. 1,920 m. 30, 32°19729"N; 106°33/517W. EV SEZ. NEA, Sec (dip smomeiee ete Upper slope of short canyon entering Ice Cn. from S. Narrow slides of rhyolitic talus in groves of Gambel Oak, interspersed with grassy areas. Probably at or near the type-locality of A. organensis. 2,225-2,255 m. 31. 32°17'42’N; 106°33’46’W. Along section line between Secs. 19 and 20, T. 23 S, R. 4 E and .6 km S of N section line. Rhyolitic talus below massive, rounded peak on south-facing wall of Soledad Cn. 1,920 m. 32. 32°14’52’N; 106°33’33’W. SW%,NE%,SW%, Sec. 5, T. 24S) Rf. 4 E. Rhyolitic talus high up on northeast-facing wall of Boulder Cn. 1,966 m. 33. 32°14717”"N; 106°33’48’W. Along section line between Secs. 7 and 8, T. 24S, R. 4 E., 48 km S of N section line. Large accumulation of rhyo- litic talus on northeast-facing wall of Finley Cn., directly opposite mouth of Long Cn. 1,623 m. Specimens of A. organensis were also taken from rhyolitic talus .5 km SW, .5 km NNW (in Long Cn.) and .8 km NE (upper end of Finley Cn.) or Ibo, 3B. Franklin Mountains, E] Paso County, Texas (not surveyed to township and. sections): 34, 31°59’51’”N; 106°30’45’”W. Rubble of limestone of Mississippian age, immediately W of “saddle” and 30 m above it; 1.2 km SE of Anthony Gap. 1,524 m. 35. 31°59’7’N; 106°30'34”’W. Talus of limestone of Mississippian age, high up on north-facing canyon wall, 2.8 km S of Anthony Gap. Canyon debouches NE into Hueco Bolson. 1,584 m. 36. 31°58’1”N; 106°30’45’W. Mouth of Vinton Cn., immediately E of a salient outcrop of massive La Tuna Limestone (Pennsylvanian), which pro- duces a “gateway” in mouth of canyon. From talus derived from the La Tuna Limestone, on north-facing wall of canyon and only slightly above arroyo floodplain in shadow of cliffs. 1,494 m. Ashmunella p. pasonis has been taken at several other places in Vinton Cn., from whence Drake (1951) obtained the type. 37. 31°57’25’N; 106°30'30”W. North-facing wall of first canyon south of Vinton Cn.; .97 km WSW of Anthony’s Nose; from talus of Mississippian limestone. 1,585 m. 38. 31°57’/18’”N; 106°29'52”W. Elongate mound of limestone talus at VOLUME 90, NUMBER 4 875 head of an east-draining canyon on E side of Franklin Mts., .4 km south of Anthony's Nose. 1,920 m. 39. 31°56’338’N; 106°30/23’W. North-facing wall of SE branch (of 3 branches) of second canyon S of Vinton Cn., on steep slope immediately north of a “saddle” connecting this canyon with canyon noted in next lo- cality. Talus of Mississippian limestone. 1,600 m. 40. 31°56’3’N; 106°30/14”W. Rubble of Silurian (Fusselman) limestone at upper, SE end of first canyon N of Tom Mays Park. 1,630 m. Al. 31°54’30-31”N; 106°30’2”W. Rhyolitic talus accumulation in can- yon indenting N side of North Franklin Mt. and debouching NW into Tom Mays Park. 1,825-1,860 m. There are numerous accumulations of rhyolitic talus and at least one of sandstone talus on the northern slopes of North Franklin Mt. All such accumulations inspected yielded A. pasonis polygyroidea. 42. 31°53’39’"N; 106°2854”W. Rhyolitic talus on south-facing slope of NW arm of Fusselman Cn., near a spring. A grove of trees near spring is readily identifiable from the trans-mountain highway, which crosses the canyon farther to the SE. 1,554 m. Acknowledgments For permitting access for collecting on White Sands Missile Range we are indebted to authorities of the Range and especially to Dr. Richard H. Duncan and Mr. B. H. Ferdig. Mr. John H. Kiger, manager, San Andres Wildlife Refuge, acted as our well-informed guide on the Refuge on two oc- casions. The invaluable contributions of Dr. Thomas K. Todsen have been mentioned above in the description of Ashmunella todseni. Literature Cited Burch, J. B. 1962. How to know the eastern land snails. Wm. C. Brown Co., Dubuque, Iowa. 214 pp. Clapp, G. H. 1908. New land shells from Arizona and New Mexico. Nautilus 22: 76-78. Cockerell, T. D. A. 1897. (Untitled). Nautilus 11:69. Drake, R. J. 1951. A new species of Polygyra from West Texas. Rev. Soc. Malacol. “Carlos de la Torre” 8:44—46. 1952. Change in genus for Polygyra pasonis. Ibid. 9:30. Mayr, E. 1963. Animal species and evolution. Harvard University (Belknap) Press, Cambridge, Massachusetts. xiv -+- 797 pp. Metcalf, A. L. 1969. Quaternary surfaces, sediments, and mollusks: southern Mesilla Valley, New Mexico and Texas, pp. 158-164. In D. A. Cordoba, S. A. Wengerd, and J. Shomaker, (eds.), Guidebook of the Border Region. New Mexico Geological! Society, Twentieth Field Conference—October 23, 24, and 25, 1969. —. 1973. New fossil Ashmunellas from New Mexico (Gastropoda: Pulmonata: Polygyridae). Veliger 16:31-39. 876 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Metcalf, A. L., and P. A. Hurley. 1971. A new Ashmunella (Polygyridae) from Dona Ana County, New Mexico. Nautilus 84(4):120-127. Metcalf, A. L., and W. E. Johnson. 1971. Gastropods of the Franklin Mountains, El Paso County, Texas. Southwestern Naturalist 16:85-109. Pilsbry, H. A. 1905. Mollusca of the southwestern states, I: Urocoptidae; Helicidae of Arizona and New Mexico. Proc. Acad. Nat. Sci. Philadelphia 57:211—290. 1915. Mollusca of the southwestern states, VI: The Hacheta Grande, Florida, and Peloncillo Mountains, New Mexico. Proc. Acad. Nat. Sci. Phila- delphia 67:323-350, pls. 5-7. —. 1936. Land shells from Texas and New Mexico. Nautilus 49:100-102. —. 1940. Land Mollusca of North America (north of Mexico). Acad. Nat. Sci. Philadelphia Monogr. 3(Vol. I, Pt. 2):vi + 575-994 + i-ix pp. Stern, E. M. 1973. The Ashmunella rhyssa (Dall) complex (Gastropoda: Polygyri- dae): Sierra Blanca-Sacramento Mountains, New Mexico. Univ. Texas at El RasOPSCiy SElg Os Vili = LOuapps Vagvolgyi, J. 1974. Eight new Ashmunellas from the southwestern United States (Pulmonata: Polygyridae). Proc. Biol. Soc. Washington 87:139-166. Van der Schalie, H., L. L. Getz, and B. C. Dazo. 1962. Hybrids between American Pomatiopsis and Oriental Oncomelania snails. Amer. Jour. Trop. Med. Hyg. 11:418—420. Webb, G. R. 1977. A laboratory hybridization study of Ashmunella (Stylommatophora, Polygyridae, Ashmunellinae). Gastropodia 1:98-99. PROC. BIOL. SOC. WASH. 90(4), 1977, pp. 877-883 A NEW SPECIES OF SYNCHELIDIUM (CRUSTACEA, AMPHIPODA) FROM SAND BEACHES IN CALIFORNIA J. Laurens Barnard Abstract—Barnard, J. Laurens, Department of Invertebrate Zoology, Smithsonian Institution, Washington, D.C. 20560.—A commonly en- countered sand beach crustacean in California is the following new species of Synchelidium. It is easily distinguished from deeper water species in the minute uropod 3, small epimeron 3 of females and the diffuse omma- tidial eyes lacking a capsule. My appreciation is extended to the several collectors of this species noted in the text. Carolyn L. Cox kindly inked and prepared the plates for publication. Synchelidium micropleon, new species Figs. 1-4 Synchelidium n. sp. Enright, 1960:758—760. Diagnosis.—Rostrum partly downturned. Eyes without distinct capsule, ommatidia scattered. Peduncle of antenna | short and stout. Right lacinia mobilis bifid. Inner plates of lower lip fully fused, lacking notch. Outer plate of maxilla 2 normally rounded, not broadened, evenly setose. Inner plates of maxillipeds narrow and elongate, bearing only 2 long apical setae. Anteroventral bevel of coxa 1 weak. Article 2 of gnathopods 1-2 densely setose anteriorly, setae mixed short and medium. Lobe of article 5 on enathopod | about 1.1 times as long as article 6; posterior margin of article 6 about 0.7 times as long as anterior margin; angle of palm 45° to longi- tudinal axis of article 6; serrations of palm of medium size. Article 6 of gnathopod 2 about 7.5 times as long as wide, not tapering distally; length of article 7 about 20% of article 6. Pereopods 3-4 of stout form, dactyls vestigial; other pereopodal dactyls of medium size. Article 2 of pereopods 5-7 poorly setose or setulose poste- riorly, each with mediofacial row of long setae; article 5 of pereopods 5-6 with anterior comb of stiff spines. Epimeron 3 of both sexes very small, narrower than epimeron 2, posterior margin sloping anteriad; epimeron 2 with several facial setae in vertical row, lower posterior margin weakly concave, posteroventral corner lacking tooth but weakly extended and rounded. Uropod 3 very small, reaching less than halfway along rami of uropod 1, peduncle very short. Telson al- most perfectly ovate. Description.—Primary flagellum of antenna 1 with 9 articles in female, 8 in male, one aesthetase each on articles 4-8 in female, upwards trom 5 on articles 1-6 in male, example of formula in male = 6—6-6—6-—7-S-l-0, 878 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON . WS Fig. 1. Synchelidium micropleon, new species, holotype female “a” 3.37 mm; h = female “h” 2.78 mm; m = female “m” 2.85 mm. A, Antenna; B, Prebuccal, anterior; C, Coxa; D, Dactyl; E, Epimeron; F, Body; G, Gnathopod; H, Head; I, Inner Plate or ramus; L, Lower lip; M, Mandible; N, Pleon; O, Outer plate or ramus; P, Pereopod; R, Uropod; S, Maxilliped; T, Telson; X, Maxilla; e, Left; 0, Apex omitted; r, Right side; s, Setae omitted. proximal to distal (density reduced in illustration); accessory flagellum ab- sent. One simple gill each on coxae 2-6, gill of coxa 6 adze-shaped, that of coxa 5 clavate, that of coxa 4 weakly clavate; others slender, sausage shaped, gill of coxa 5 dominant. Brood plates long, slender, with long setae mostly apical, only 3 pairs present, thus absent on coxa 2. VOLUME 90, NUMBER 4 879 be a \ Ay, D3 Fig. 2. Synchelidium micropleon, new species, holotype female “a” 3.37 mm; m female “m” 2.85 mm; x = male “x” 2.39 mm. See Fig. 1 for symbols. 880 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 3. Synchelidium micropleon, new species, holotype female “a” 3.37 mm; m = female “m” 2.85 mm. See Fig. 1 for symbols. VOLUME 90, NUMBER 4 881 \- Fig. 4. Synchelidium micropleon, new species, holotype female “a” 3.37 mm; m female “m” 2.85 mm. See Fig. 1 for symbols. 882 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Spination on rami of uropods 1-3 variable in voucher material, usually outer ramus of uropod 1 with 3 spines, inner with 2, outer of uropod 2 with 2 spines, inner with 2, outer of uropod 3 with 2, inner with one; occasion- ally outer ramus of uropod 1 with 5 spines, outer of uropod 2 with 4, inner with 3, or in smaller specimens rami of uropod 3 each with one spine and inner of uropod 2 with one spine. Medial apex of peduncle on uropod 2 with one spine slightly larger than that illustrated for lateral apex. Illustration of detached uropod 1 reduced in magnification from that of uropod 2. No fully terminal male available in collections, illustrated male with slightly elongate flagellum of antenna 2; presumably terminal males swim in neritic nekton and will have to be captured in plankton tows. Relationship.—This species differs from all known species in the very small uropod 3 which reaches less than halfway along the rami of uropod 1. In addition, epimeron 3 of the female is narrowed as in males of various species; as far as can be determined epimeron 3 on females of other species is as broad as or broader than epimeron 2. The known species of Synchelidium are outlined below and, for each, one or more additional characters of difference from S. micropleon are stated: Synchelidium haplocheles (Grube) (See Sars, 1895, as S. brevicarpum; Stebbing, 1906): longer more slender peduncles of antennae in female, much broader inner plate of maxilliped with shorter, more numerous setae; subrectangular, emarginate telson. Northeastern Atlantic-Mediterranean. Synchelidium intermedium Sars (1895): short lobe on wrist of gnathopod 1, non-ovate telson. Northeastern Atlantic-Mediterranean. Synchelidium tenuimanum Norman (see Sars 1895, as S. haplocheles): short wrist of gnathopod 1, non-ovate telson. Northeastern Atlantic. Synchelidium maculatum Stebbing (1906) (see Chevreux and Fage, 1925): short wrist of gnathopod 1, non-ovate telson. Northeastern Atlantic- Mediterranean. Synchelidium longidigitatum Ruffo (1947): enlarged dactyls of pereopods 3-4; many more posterior setae on article 2 of pereopods 5-7. Mediter- ranean. Synchelidium shoemakeri Mills (1962): sharper tooth on epimeron 2, non-ovate telson, more numerous posterior setae or setules on article 2 of pereopods 5-7, broader inner plates of maxilliped with more numerous shorter setae. Northeastern Pacific. Synchelidium rectipalmum Mills (1962): epimeron 2 lacking postero- ventral excavation, non-ovate telson, more numerous posterior setae on article 2 of pereopods 5-7, broader inner plates of maxillipeds with more numerous shorter setae, palm of gnathopod 1 transverse, dactyl of gnatho- pod 2 much longer, hand shorter. Northeastern Pacific. VOLUME 90, NUMBER 4 883 Synchelidium miraculum Imbach (1969): dactyls of pereopods 3-4 large, female antennal peduncles much more elongate, article 2 of antenna 1 especially elongate, article 3 short, short wrist of gnathopod 1. South China Sea. Synchelidium americanum Bousfield (1973): less oblique palm of gnathopod 1, non-ovate telson, unexcavate epimeron 2, broader inner plates of maxillipeds with more numerous shorter setae, deeply notched inner lobes of lower lip, stronger posterior setation on article 2 of pereopods 5-7. Northwestern Atlantic. Holotype.—USNM No. 109895, female “a” 3.37 mm (illustrated). Type-locality—San Francisco Ocean Beach, California, 13 September 1959; intertidal, J. T. Enright. Paratypes.—Type-locality, female “a” 3.37 mm (illustrated), female “h” 2.78 mm (illustrated), female “m” 2.85 mm (illustrated), female “p” 2.32 mm and 4 other specimens. Voucher material: Morro Bay, California, 9 Sep- tember 1959, intertidal, J. T. Enright, male “x” 2.39 mm (illustrated) and 76 other specimens. Other material—Dillon Beach, Marin County, California, 19 September 1959, intertidal, J. T. Enright (2); La Jolla California, beach in front of Scripps Institution of Oceanography, intertidal, following collections: Miarehy 1938; net, Olga Hartman (13); 1957-1958, 6 m, E. W. Fager (25); 15 May 1958, J. T. Enright (100+); Estero de Punta Banda, Baja Cali- fornia, 23 February 1952, sandy beach, T. E. Bowman (2). Distribution.—Dillon Beach, California to Estero de Punta Banda, Mex- ico, 0-6 m, mainly on sand beaches. Literature Cited Bousfield, E. L. 1973. Shallow-water Gammaridean Amphipoda of New England. Comstock Publishing Associates, London & Ithaca. 312 pp., 69 pls. Chevreux, E., and L. Fage. 1925. Amphipodes. Faune de France. 9:1—488, figs. 1-438. Enright, J. T. 1960. Pressure sensitivity of an amphipod. Science 133(3455): 760. Imbach, M. C. 1969. Gammaridean Amphipoda from the South China Sea. Naga Report 4:39-167, 33 pls. Mills, E. L. 1962. Amphipod crustaceans of the Pacific coast of Canada, UH. Family Oedicerotidae. Natural History Papers National Museum Canada 15:1—21, 6 figs. Ruffo, S. 1947. Studi sui Crostacei Anfipodi XII Gli Anfipodi del Museo Civico di Storia Naturale di Genova C) Su aleuni Anfipodi Mediterranei e descrizione di una nuova species del gen. Synchelidium G. O. Sars. Annales Museo Civico Storia Naturale Genova 63:79-89, 2 figs. Sars, G. O. 1895. Amphipoda. An account of the Crustacea of Norway with short descriptions and figures of all the species, l:i-viii + 1-711, pls. 1-240, supple- mentary pls. 1-8. Stebbing, T. R. R. 1906. Amphipoda I. Gammaridea. Das Tierreich, 29:1—S06, figs 1-197. 5s- PROC. BIOL. SOC. WASH. 90(4), 1977, pp. 884-893 STILBOMASTAX, A NEW GENUS OF SPIDER CRAB (MAJIDAE: TYCHINAE) FROM THE WEST INDIES REGION, WITH NOTES ON AMERICAN RELATIVES! Austin B. Williams, J. Kevin Shaw, and Thomas S. Hopkins Abstract.—Williams, A. B., National Marine Fisheries Service System- atics Laboratory, National Museum of Natural History, Washington, D.C., 20560, J. K. Shaw, and T. S. Hopkins, Dauphin Island Sea Lab, P.O. Box 386, Dauphin Island, Ala. 36528.—A new genus, Stilbomastax, is recog- nized from the West Indies and Gulf of Mexico for reception of the spider crab Tyche margaritifera Monod, 1939 (= Stilbognathus burryi Garth 1952). The new genus in some respects lies between Tyche from the western hemisphere and Stilbognathus from the western Indian Ocean, and comparisons are made among species belonging to these genera. Monod (1939) described a unique male spider crab from Guadeloupe as Tyche margaritifera. Garth (1952a), unaware of Monod’s paper, described a similar “Tyche-like” female from southeast Florida as Stilbognathus burryi, believing this to be one of the interesting rare extensions of essen- tially Indo-Pacific genera into the Western Atlantic; but he also stated (p. 251) that “the erection of a new genus might be justified,” basing his reasoning on Balss’s (1929) distinction of Tyche from Stilbognathus on the basis of free vs. fused female abdominal segments. Guinot (1964), review- ing the related genera Tyche Bell 1835 (Western Atlantic-Eastern Pacific), Stilbognathus von Martens 1866 (Red Sea and east Africa), and Ophthal- mias Rathbun 1897 (western Indian Ocean), left the generic placement of T. margaritifera and S. burryi uncertain, although she thought that they were possibly synonymous within the genus Tyche. She had only Monod’s specimen and Garth's (1952a) figures for study. Mature male and female specimens recently collected in the eastern Gulf of Mexico provide ample evidence that Monod’s and Garth’s species are synonymous, as well as evi- dence to support Garth’s idea of generic independence. Stilbomastax, new genus Postorbital lobe forming commencing orbit. Mouth frame (Fig. 1) di- vergent anteriorly, flared and thickened into strong rim at anterolateral cor- ners; anterior margin thin and raised. External maxillipeds with merus deeply inserted into outer border of ischium, its central part porcellanous, hemispheric, but with thin alate mesial and anterolateral lobes; ischium deeply and broadly grooved longitudinally, its thin distal edge closely covering proximal part of meral swelling; a less prominent longitudinal groove between outer border of endognath and excavate inner border of VOLUME 90, NUMBER 4 885 exognath; curved or straight prolongation at base of exognath overlapping ischium of endognath ventrally. Female abdomen (Fig. 8) almost circular in outline, segments 4-6 fused but evident. Male abdomen with 7 segments free, ribbed and uneven; first pleopods (Fig. 10a) of simple, somewhat flattened form distally, with short tip bent abruptly laterad and slightly re- flexed to lateral opening. Type-species.—Tyche margaritifera Monod 1939. Etymology.—From the Greek “stilbo” glitter or gleam, and “mastax” jaws. The gender is feminine. Remarks.—Tyche has an essentially rectangular mouth frame whose rim is variably raised at its anterolateral corner, depending on the species. Both Ophthalmias and Stilbognathus have a mouth frame with sides obviously diverging anteriorly; in Ophthalmias the ischium of the external maxillipeds is notched along its posteromesial margin, adjacent to the intervening tri- angular sternal plate, while the corresponding surface in Stilbognathus lacks this notch (Guinot 1964). In these characters Stilbomastax stands be- tween Tyche and Stilbognathus, having a mouth frame less divergent ante- riorly than the latter. Species of these genera may have one or all of the following features on the external maxillipeds: porcellanous surfaces on ischium or merus, swollen in some species; longitudinally grooved ischium; lamellar distal expansion on ischium covering merus proximally except along lateral side at articulation. Only Tyche and Stilbomastax have a basal prolongation on the exognath of the external maxillipeds (although there is a rudimentary projection in Stilbognathus erythraeus yon Martens), hence in the last couplet of Garth’s (1958a:162) Key to New World genera of the subfamily Ophthalmiinae Stilbomastax should be substituted for Stilbognathus. Abdominal segments of female Tyche are free. Segments 4-6 are fused in Stilbognathus and Stilbomastax but in the former the segments are so united that they are nearly obliterated whereas in Stilbomastax margariti- fera the segments are individually raised and easily recognizable. Male first pleopods of Tyche and Stilbomastax are similar (Fig. 10a, b; also Garth 1958b:pl. J, figs. 4-6), being least bent and reflexed distally in T. emarginata White. In Stilbognathus species the tip is flattened, differently bent and membranously ornamented (Fig. 10c). The characters found in Stilbomastax margaritifera that are shared with Tyche and Stilbognathus but differently combined, along with indepen- dent ones, are best accommodated by placing this species in a distinct genus. Notes on Some American Representatives of the Subfamily Tychinae Tychinae Tychinae Dana 1851:43; (including Tyche Bell). Stenocionopinae Miers 1879:652 (part: including Stilbognathus von Mar- tens and Tyche Bell). 886 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON VOLUME 90, NUMBER 4 887 Stenociopoida Alcock 1895:161, 166 (part: including Stilbognathus von Martens and Tyche Bell). Ophthalmiinae Balss 1931:6 (name substituted for Stenocionopinae Miers). —Sakai 1938:243.—1976:187.—Garth 1958a:161 (redefined).—Williams 1965:246.—Griffin 1966:262, 263, fig. 1 (sensu Garth 1958a). Both Rathbun (1925) and Balss (1957) included genera discussed here in the subfamily Majinae Alcock 1895 (broad sense). The following list contains citations of original descriptions, principal references, type-localties, distribution records, and emended descriptions of the mouthparts for all species of Tyche as well as Stilbomastax margar- itifera, with selected measurements for the latter. Stilbomastax margaritifera (Monod) Figs) 1-3, 10a Tyche margaritifera Monod 1939:561, figs. 6, 7, 8, 9—Guinot 1964:45, fig. Be Tol, 45, inks JE Stilbognathus burryi Garth 1952a:252, pl. 1—Guinot 1964:45, 51-53 (here and there). Type-locality— Basse-Terre, Guadeloupe, 15-20 m. Known range.—SE of Cape San Blas to SE Florida; Guadeloupe. Habitat —Garth (1952) noted the habitat as hard rock, broken shell, 38 m. Specimens from west Florida were found on hard rock and coral shell rubble among sponges, corals, and algae. Mouthparts.—Mouth frame with thin anterior margin moderately raised, anterolateral margin thickened into strong rim. External maxillipeds with ischium broadly and longitudinally grooved centrally, tooth on mesial margin and with oblique, thin, flared anterior lobe covering edge of swollen merus; merus with deep proximal insertion along lateral margin of ischium, central part a glistening white hemisphere, anterolateral blade thin, tri- angular, projecting, and thin anteromesial margin trilobate; exognath with inner border excavate, basal prolongation variable, projecting either straight posteromesially or curving mesiad, sometimes laterad, spur on protopodite lateral to base of prolongation rather broad. New material—USNM 168479, 45 mi W Sanibel Island, Fla., 26°25’N, 82°97'32” W, 36.6 m, 28 June 1976; mature ¢, carapace length 27.8 mm, < Figs. 1-6. External maxilliped: 1. Stilbomastax margaritifera (USNM 168480); 2. Tyche potiguara (after Garth 1952b); 3. T. lamellifrons (USNM 72668); 4. T. emarginate (after Garth 1946); 5. T. clarionensis (after Garth 1958a); 6a. T. galapagensis (USNM 100917); b, punctate surface magnified. Scales: 1-Ga = 1 mm, Gh 5 mm 888 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON a4 as Ses y SS PENSE aS Figs. 7-9. Mature female abdomen: 7. Tyche emarginata (USNM 46770); 8. Stilbo- mastax margaritifera (USNM 168480); 9. Stilbognathus erythraeus (USNM 169304). 10. Male first pleopod: a, Stilbomastax margaritifera (USNM 168479); b, Tyche emarginata (after Williams 1965); c, Stilbognathus erythraeus (after Guinot 1964, fig. 37b). Scales: 1-9 — emma — eon aby oonmm width 19.5; rostrum | 4.38, w 3.9, exorbital w 13.2, hepatic constriction w 12.5, posterior notch to gastric summit | 14.5, right cheliped palm height 4.0, 1 9.5, dactyl | 4.6, ischio-merus | 9.38; R/V Bellows, trawl, T. S. Hop- kins. USNM 168480, Florida Middle Ground, 80 mi W Tarpon Springs, Fla., 28°38’N, 84°19’W, 33 m, 11 June 1974; 2 ov, carapace | 28.8, w 21.5; R/V Bellows, SCUBA, T. W. Hopkins. VOLUME 90, NUMBER 4 889 Dauphin Island Sea Lab: MAFLA-II-N, Florida Middle Ground, 80 mi W Tarpon Springs, Fla., 28°24’N, 84°21’W, 36 m, 11 June 1974; é carapace | 25.4, w 17.4; R/V Bellows, dredge, T. S. Hopkins. 33-194-IV-A-a, about 35 mi W Cape San Blas, Fla., 29°04’N, 85°14’W, 36.6 m, 26 February 1976; 2 carapace | 25.2, w 18.3; R/V Bellows, dredge, T. S. Hopkins. Florida Department of Natural Resources: 36 mi W Egmont Key, Fla., 27°37'N, 83°28’W, 37 m, R/V H. Cortez, B. Presley. FSBC I 1548, trawl, 3 January 1966; 2¢, carapace | 23.5-26.3, w 15.0-16.7. FSBC I 17511, dredge, 11 August 1967; 6 carapace | 27.8, w 19.0. FBSC I 17512, dredge, 6 January 1967; ¢ carapace | 35.0, w 24.8. FSBC I 17513, dredge, 20 May, 1967; 23, carapace | of one 13.9, w 9.4. Tyche potiguara Garth Higw2 Tyche potiguara Garth 1952b:45, figs. 1-8.—Coelho and Ramos 1972:210. Type-locality Off Cabo de Sao Roque, Brazil, 06°59’30’S, 34°47’W, 36.6 m, Albatross 2758. Known range.—Paraiba to Alagoas, Brazil. Mouthparts—Mouth frame with anterior margin slightly raised, antero- lateral margin slightly thickened. External maxillipeds with ischium shal- lowly grooved longitudinally, toothed on mesial margin and with anterior lobe covering merus proximally; merus with porcellanous, convexly menis- coid anteromesial lobe separated from narrow, elongate anterolateral lobe by longitudinal depression, body of merus inserted deeply into ischium laterally; exognath with strong somewhat flattened prolongation recurved to lie in proximal part of ischial groove, spur on protopodite lateral to base of prolongation. Tyche lamellifrons Bell Fig. 3 Tyche lamellifrons Bell 1835:173.—Garth 1958a:173.—1958b:pl. J, fig. 4: pl. 18, fig. 2 (not Garth 1946:406, the Galapagos specimens). Type-locality —Panama. Known range.—Agua Verde Bay, Gulf of California, Mexico, to La Libertad, Ecuador (not Galapagos Islands, as reported by Garth (1946)). Mouthparts—Mouth frame with anterior margin moderately raised, an- terolateral margin moderately raised and thickened. External maxillipeds with ischium broadly and longitudinally grooved, toothed on mesial mar- gin, and with an anteromesial expansion covering merus proximally; merus with anteroexternal angle squared or rounded off, not extended as thin blade, mesial aspect trilobate; exognath with basal prolongation recurved 890 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON to lie in proximolateral entrance to central groove of ischium, spur on — protopodite lateral to base of prolongation. Tyche emarginata White Figs. 4, 10b Tyche emarginata White 1847:206.—Rathbun 1925:508, pl. 272; pl. 273, figs. 7-12.—Garth 1946:408-408 (here and there), text-fig. 1—Williams 1965:247, figs. 225, 226, 233B. Type-locality.—West Indies. Known range.—Off Beaufort Inlet, N.C., through Bahamas to west coast of Florida. Mouthparts—Mouth frame with anterior margin slightly and antero- lateral corner moderately raised and thickened. External maxillipeds smooth, shining and slightly inflated; ischium strongly toothed mesially, its broad laterally expanded anterior lobe overlying merus proximally; promi- nent merus deeply inserted into distolateral corner of ischium, with rounded thin lobe at anterolateral corner and internal corner divided into 3 narrowly separated lobes; exognath with large basal prolongation re- curved to fit flush into shallow groove on base of ischium, spur on pro- topodite lateral to base of prolongation. Tyche clarionensis Garth Fig. 5 Tyche clarionensis Garth 1958a:176, fig. 5.—1958b:pl. J, fig. 5. Type-locality Sulphur Bay, Clarion Island, Mexico, 55 m. Known range.—Clarion Island. Mouthparts—Endognath of outer maxilliped smooth and_ inflated; ischium toothed along mesial margin, lateral expanded anterior lobe covering merus proximally; merus inserting deeply into ischium laterally, anterolateral angle produced into thin blade, anteromesial margin cut into 3 distinct lobes; hooked prolongation of exognath lodged in basal groove of ischium, completely filling it, spur on protopodite lateral to base of pro- longation. Tyche galapagensis Garth lene, (6 Tyche galapagensis Garth 1958a:178.—1958b:pl. J, fig. 6. Type-locality—Albemarle Point, Albemarle Island, Galapagos Islands, shore. Known range.—Galapagos Islands: also Sullivan Bay, James Island; VOLUME 90, NUMBER 4 891 Post Office Bay, Charles Island; Marchena Island; west of Gardner Island, Hood Island; Darwin Bay, Tower Island (see also Garth 1946:406). Mouthparts—Mouth frame with low anterior margin and slightly thick- ened anterolateral corner. External maxillipeds rather broad, margins hairy; endognath with ischium and merus inflated, smoothly convex but densely and finely punctate, superficially seeming almost coalesced; ischium with roughly rectangular anteromesial lobe overlapping part of merus, mesial margin with irregular teeth hidden in hairs; merus inserted deeply into outer margin of ischium, anteriorly narrowing thin and blade- like with distal margin serrate, mesial margin trilobate; exognath with basal prolongation curved but directed posteromesially rather than recurving onto ischium, spur on protopodite lateral to base of prolongation remote. Key to Species Based on Endognath of External Maxillipeds bo 1. Ischium broadly and deeply grooved longitudinally — Ischium not broadly and deeply grooved longitudinally 3 2. Merus swollen to glistening white hemisphere centrally Stilbomastax margaritifera. — Merus not swollen centrally Tyche lamellifrons. 3. Ischium-merus smoothly convex, appearing fused 4 — Ischium-merus neither smoothly convex nor appearing fused 5 4. Ischium-merus smooth and shining; mesial teeth easily visible T. emarginata. — Ischium-merus not shiny but uniformly punctate; mesial teeth hidden in hairs T. galapagensis. 5. Unswollen merus with margin trilobate near palp T. clarionensis. — Somewhat swollen % of merus with margin entire near palp T. potiguara. Acknowledgments We thank John S. Garth for generously discussing and critically read- ing the manuscript. We also thank J. Forest, Muséum National d Histoire Naturelle Paris, L. B. Holthuis, Rijksmuseum van Natuurlijke Historie, Leiden, and David K. Camp, Florida Department of Natural Resources, St. Petersburg, for loan of study material. Initial field work was supported by a subcontract of the Bureau of Land Management Contract No. $550-CTS- 30 to T. S. Hopkins. The illustrations were prepared by Maria M. Dieguez. Literature Cited Alcock, A. W. 1895. Materials for a carcinological fauna of India. No. 1. The Brachyura Oxyrhyncha. Journal of the Asiatic Society of Bengal 64: 157-291, pls. 3-5. 892 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Balss, H. 1929. Decapoden des Roten Meeres. IV. Oxyrhyncha und Schlussbetrach- tungen. Denkschriften Akademie der Wissenschaften, Wien, Mathematisch- Naturwissenschaftliche Klasse 102: 1-30. —. 1957. Decapoda. VIII. Systematik. Bronns Klassen und Ordnungen des Tierreichs. Bd. 5, Abt. I, Buch 7. Lf. 12:1050-1672. Bell, T. 1835. Some account of the Crustacea of the coasts of South America, with descriptions of new genera and species; founded principally on the collections obtained by Mr. Cuming and Mr. Miller. (Tribus 1, Oxyrhynchi). Proceedings of the Zoological Society of London 3:169-173. 1836. Some account of the Crustacea of the coasts of South America, with descriptions of new genera and species; founded principally on the collections obtained by Mr. Cuming and M. Miller. Transactions of the Zoological Society of London 2:39-66, pls. 8-13. Coelho, P. A., and M. de A. Ramos. 1972. A constituicdo e a distribuicao da fauna de decapodos do littoral leste de America do Sul entre as latitudes de 5°N e 39°S. Trabalhos do Instituto Oceanografia Universidade Federal Recife 13:133-236. Dana, J. D. 1851. On the classification of the maioid Crustacea or Oxyrhyncha. American Joumal of Science and Arts, Ser. 2, 11:425-434. Desbonne, I. 1867. In I. Desbonne and A. Schramm, Crustacés de la Guadeloupe d’aprés un manuscrit du... I. Desbonne comparé avec les échantillons de crustacés de sa collection et les derniéres publications de MM. H. de Saussure et W. Stimpson. I. Partie. Brachyures. [Edited, with a preface, by A. Schramm] 60 pp., pls. 1-8. Garth, J. S. 1946. Littoral brachyuran fauna of the Galapagos Archipelago. Allan Hancock Pacific Expeditions 5(10):i-iv, 341-601, pls. 49-87. 1952a. A review of the crustacean genus Stilbognathus von Martens (Deca- poda, Maiidae) with description of a new species from the east coast of Florida. Bulletin of Marine Science of the Gulf and Caribbean 1(4):249-256, 1 pl. ——. 1952b. “Tyche potiguara,’ a new species of decapod crustacean from Brazil (Maiidae, Ophthalmiinae). Revista Brasiliera de Biologia 12(1):45—48. ——. 1958a. Brachyura of the Pacific coast of America, Oxyrhyncha. Allan Han- cock Pacific Expeditions 21(1):xii + 499 pp. 1958b. Brachyura of the Pacific coast of America, Oxyrhyncha. Tables and plates. Ibid. 21(2):501—854, pls. A-Z, 1-55. Griffin, D. J. G. 1966. A review of the Australian majid spider crabs (Crustacea, Brachyura). Australian Zoologist 13(3):259-298, pls. 15-17. Guinot, D. 1964. Sur une collection de crustacés décapodes brachyoures de Mer Rouge et de Somalie. Remarques sur les genres Calappa Weber, Menaethiops Alcock, Tyche Bell, Ophthalmais Rathbun et Stilbognathus von Martens. Bol- lettino del Museo Civico di Storia Naturale di Venezia 15 (for 1962):7—63, pls. 1-4. Martens, E. von. 1866. Verzeichniss der von Dr. E. Schweinfurth im Sommer 1864 auf seiner Reise am Rothen Meere Gesammelten und nach Berlin eingesendeten zoologischen Gegenstande. II. Crustaceen. Verhandlungen der Zoologisch- Botanischen Gesellschaft in Wien 16:377-382. Miers, E. J. 1879. On the classification of the maioid Crustacea or Oxyrhyncha, with a synopsis of the families, subfamilies, and genera. Jourmal of the Linnean Society, London, Zoology 14:634—673, pls. 12-13. Monod, T. 1939. Sur quelques crustacés de la Guadeloupe (Mission P. Allorge, 1936). Bulletin du Museum National d’Histoire Naturelle, Paris. Ser. 2, 11(6): 557-568. - ——____—- a VOLUME 90, NUMBER 4 893 Rathbun, M. J. 1925. The spider crabs of America. United States National Museum Bulletin 129:xx + 613 pp., 283 pls. Sakai, T. 1938. Studies on the crabs of Japan. III. Brachygnatha, Oxyrhyncha. Yo- kendo, Tokyo, pp. 193-364, pls. 20-41. 1976. Crabs of Japan and the adjacent seas. Kodansha Ltd., Tokyo, 773 pp. [English text]. White, A. 1847. Short descriptions of some new species of Crustacea in the collec- tion of the British Museum. Annals and Magazine of Natural History 20(132): 205-207. Williams, A. B. 1965. Marine decapod crustaceans of the Carolinas. United States Fish and Wildlife Service Fishery Bulletin 65(1):xi + 298 pp. Footnote 1 Contribution from National Marine Fisheries Service Systematics Laboratory, and No. 011 from Marine Environmental Sciences Consortium, Dauphin Island, Ala. PROC. BIOL. SOC. WASH. 90(4), 1977, pp. 894-914 CYCLOLEBERIS CHRISTIEI, A NEW SPECIES OF MARINE OSTRACODA (SUBORDER MYODOCOPINA) FROM SALDANHA BAY AND LANGEBAAN LAGOON, SOUTH AFRICA Louis S. Kornicker and Rosalie F. Maddocks Abstract.—Kornicker, L. $., Department of Invertebrate Zoology, Smith- sonian Institution, Washington, D.C. 20560, and R. F. Maddocks, Depart- ment of Geology, University of Houston, Houston, Texas 77004.— Cycloleberis christiei, a new species of marine Ostracoda (Suborder Myodocopina) from Saldanha Bay and Langebaan Lagoon, South Africa, is described and illustrated, and the ontogeny and variability of the species is discussed. Saldanha Bay and Langebaan Lagoon form a connected bay-lagoonal complex about 24 km in length along the western coast of South Africa about 106 km north of Cape Town. The northern part, Saldanha Bay, is open to the Atlantic Ocean, whereas Langebaan Lagoon, a shallow lagoon, is open to Saldanha Bay at its northern end but is separated from the ocean by a narrow peninsula along its western shore. For map of the bay and lagoon see Day (1959: fig. 1). According to Day (1959:486) surface water temperatures range from 13.9°C in winter to 16.9°C in summer in Saldanha Bay, while at the south- ern end of Langebaan Lagoon the annual range is 9.9°C-24.0°C; salinities in Saldanha Bay range from 34.62-34.91%, and are as low as 32.66% in the southern part of Langebaan Lagoon after a winter rain. The specimens upon which this study is based are from individual col- lections made by Drs. John H. Day, Nigel Christie, and Anton Moldan, all of the Zoology Department, University of Cape Town, Rondebosch, South Africa. We are indebted to them for making the material available. The new species described herein brings to 3 the number of species of the genus Cycloleberis reported to be living along the western coast of Africa. In the description of this species we include information concerning its on- togeny and variability. > Fig. 1. Cycloleberis christiei, holotype USNM 157334, ovigerous female, outside views of left valve: a, Complete valve, length 5.0 mm; b, Rostrum and incisur, note attached protozoans, x50; c, Oblique ridges and fossae containing bristles on antero- ventral valve surface, 300; d, Detail from c, x 1,500; e, Scalloped ridge along anterior margin of rostrum, from b, «620; f, Anterior view of valve showing scalloped marginal ridge and bristles, 240; g, Detail from f, 540; h, Detail of pore and bristle near middle of f, x3,500. Micrographs reduced to 55% for publication. a joa lbesiiiioe ZA SO lil ~t pci a —~Q = i 5) Z, - o>) = = =) a eo) > 896 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON The holotype and paratypes of the new species have been deposited at the National Museum of Natural History, Smithsonian Institution. Five para- types have also been sent to Dr. John H. Day, University of Cape Town. We are grateful to Anne Cohen, Smithsonian Institution, for reviewing our manuscript. CYLINDROLEBERIDIDAE Miuller, 1906 CYCLASTEROPINAE Poulsen, 1965 Cycloleberis Skogsberg, 1920 Cycloleberis christiei, new species Figs. 1-7 Cycloleberis galatheae Poulsen, 1965:260, fig. 89. [juvenile ¢ paratype from Beira, Mozambique, not holotype from False Bay, South Africa. Referral is considered questionable]. Etymology.—The species is named for Dr. Nigel Christie, who supplied some of the specimens described herein. Distribution Saldanha Bay, at depths of 5-24 m in mud, fine silt, fine sand, shell and sand, and coarse shell and sand. Langebaan Lagoon, at depths ranging from intertidal to 7 m in muddy sand, silt and sand, fine sand, medium sand, sand among rocks, sand and gravel, and coarse shelly sand. Holotype —USNM 157334, ovigerous female on slides and in alcohol. Type-locality —Langebaan Lagoon, South Africa, sta. LB 601M, 17 Feb. 1975,.33°07'S, 18°01’12”E, 3.5 m, fine sand. Paratypes.—Langebaan Lagoon: Sta LB 184, 26 April 1949, 33°12’S, 18°05’E, intertidal, muddy sand: USNM 157292, 157336, 157364—1 adult é and 2 adult 2. Sta LB 218, 26 April 1949, 33°12’S, 18°05’E, intertidal, muddy sand: USNM 157347—1 adult ?. Sta LB 230, 12 Dec. 1950, 33°07’S, 18°03’E, intertidal, sand among rocks: USNM 157293, 157294, 157324—3 A-1¢é, USNM 157361—1 adult ¢. Sta LB 240, 2 May 1951, 33°07’S, 18° 03’E, depth and substrate unrecorded, probably intertidal: USNM 157295 —l A-24, USNM 157296—1 A-14, USNM 157327—1 A-36, USNM 157406 —1 A-12. Sta LB 241J, 2 May 1951, 33°07’S, 18°03’E, depth unrecorded, probably intertidal, sand among rocks: USNM 157335, 157337, 157338, 157339—4 adult 2, USNM 157350, 157358—2 A-1?, USNM 157333—1 A- 26, USNM 157328, 157329—2 A-36, USNM 157346, 157354, 157355, 157356, 157357, 157359—7 juv. 2. Sta LB 243, 2 May 1951, Sandy Point, intertidal, sand: USNM 157362—1 adult °. Sta LB 267, 3 May 1951, 33° 07'S, 18°03’E, depth and substrate unrecorded, probably intertidal: USNM 157353—1 A-12, USNM 157322—1 A-26, USNM 157360—1 juv. @. Sta 409, 6 Dec. 1953, 33°04’S, 18°02’E, intertidal, sand: USNM 157349—1 A-1 2, USNM 157297—1 A-23, USNM 157407—1¢. Sta LB 447Q, 1 May VOLUME 90, NUMBER 4 897 1954, 33°07’S, 18°03’E, depth and substrate unrecorded, probably inter- tidal: USNM 157340, 157344—2 adult ?, USNM 157331—1 A-34, USNM 157323—1 A-1¢é, USNM 157330—1 A-36, USNM 157341, 157342—2 A-12, USNM 157343—1 juv. @. Sta LB 448G, 1 May 1954, 33°07’S, 18°03’E, intertidal: USNM 157291—1 adult ¢, 157363, 157351—2 adult 2, USNM 157299, 157325—2 A-26, USNM 157326, 157332—2 A-34, USNM 157345, 157348, 157352—3 juv. Sta LB 474, 8 May 1955, 33°07’S, 18°03’E, inter- tidal, sand: USNM 157298—1 A-1é. Sta LB 577N, 15 Dec. 1974, 33°06’54” S, 18°01’48”E, 4.5 m, fine sand: USNM 157390—3 spec. Sta LB 578D, 15 Dec. 1974, 33°07’S, 18°01’12”E, 3.5 m, fine sand: USNM 157392—7 spec. Sta LB 579P, 15 Dec. 1974, 33°06’06”S, 18°01’06”E, 4.5 m, fine sand: USNM 157388—1 spec. Sta LB 582J, 20 Dec. 1974, 33°05/24”S, 18°09’E, 4.5 m, silt and sand: USNM 157386—5 spec. Sta LB 585G, 19 Dec. 1974, 33°05'12”S, 18°04’E, 7 m, sand and gravel: USNM 157391—1 spec. Sta LB 588H, 17 Dec. 1974, 33°05’24”S, 18°00’E, 2.1 m, medium sand: USNM 157387—1 spec. Sta LB 589H, 16 Dec. 1974, 33°08’30”S, 18°03’54’E, 2.1 m, fine sand: USNM 157389—1 spec. Sta LB 591E, 18 Feb. 1975, 33°05'42’S, 18°01’12”E, 3 m, silt and sand: USNM 157380—1 juv. Sta LB 593J, 18 Feb. 1975, 33°05’30”S, 18°00’04”E, 6.5 m, sand and gravel: USNM 157385— I spec. Sta LB 596G, 18 Feb. 1975, 33°05’24”S, 18°00’E, 2.1 m, medium Sand USN NM lo7394—2 spec. Sta LB S97F, 20 Feb: 1975, 33°0554’S, 18°01’24”E, 5 m, coarse shelly sand: USNM 157383—9 spec. Sta LB 598F, 17 Feb. 1975, 33°06’06”S, 18°01’06”E, 4.5 m, fine sand: USNM 157382— lspecy ora OB S99, 17 Keb. 1975, 33°06718”S, 18°0136”E, 5.8 m, medium sand: 5 specimens sent to Dr. John H. Day, University of Cape Town. Sta LB 600P, 17 Feb. 1975, 33°06’54’S, 18°01’48’E, 4.5 m, fine sand: USNM 157384—7 spec. Sta LB 601M, 17 Feb. 1975, (type-locality): USNM 157381 —46 specimens. Sta LB 602F, 17 Feb. 1975, 33°08’24”S, 18°02’06”E, 7 m, fine sand: USNM 157379—1 adult ¢. Sta LB 603F, 18 Feb. 1975, 33°08’ » 3307S, 18°03’54’E, 2.1 m, fine sand: USNM 157378—1 adult ¢. Sta LB 7 604N, 20 Feb. 1975, 33°09’30’S, 18°04’24”E, 3.1 m, fine sand: USNM ' 157396, 46 specimens. Sta LB 605J, 20 Feb. 1975, 33°09’30”S, 18°05’06” E, 2m, fine sand: USNM 157393—25 spec. Sta LB 591E, LB 602F, LB 603F (combined samples)—USNM 157290—2 adult ¢, 157318, 157319—2 adult 6, USNM 157377—3 spec. Saldanha Bay: Sta SB 295V, 30 April 1963, 33°03’54”S, 18°00’30”E, 14 m, fine sand: USNM 157401—4 spec. Sta SB 297K, 30 April 1963, 33°03'54”S, 18°00’30”E, 14 m, fine sand: USNM 157399—1 spec. Sta SB 308V, 30 April 1963, 33°03’42’”S, 18°00’49”E, 13 m, mud: USNM 157398—1 spec. Sta SB 310V, 1 May 1963, exact locality not recorded, 5 m, coarse sand and shell: USNM 157400—4 spec. Sta SB 350L, 29 April 1964, 33°02’54”S, LS°O1'S6" FE, 13 m, shell and sand: USNM 157397—1 spec. Sta SB 410F, 12 April 1975, 83°00/21”S, 17°57’'18”E, 5 m, fine sand: USNM 157395—1 juv. Sta SB 898 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 2. Cycloleberis christiei. a-d, Holotype, USNM 157334, ovigerous female: a, Complete specimen, length 5.0 mm; b, Medial eye and rod-shaped organ; c, Outline of right lateral eye; d, Upper lip, anterior to right. e, Paratype, adult female, USNM 157364, 6th limb. A1IC, 4 July 1975, 33°03’07”S, 17°58’05’E, 24 m, fine silt: USNM 157405 —I' juv. Sta SB 413C, 4 July 1975, 33°03’S, 18957547. SO eieeeeiniemesthte: USNM 157402—1 spec. Sta SB 419C, 4 July 1975, 33°00’05”S, 17°57’09”E, 9 m, fine silt: USNM 157404—1 spec. Sta SB 429B, 4 July 1975, 33°01’00"S, 18°00’00’E, 15 m, fine silt: USNM 157403, 1 juv. Description of adult female.—Carapace similar to that of adult female of Cycloleberis squamiger (Scott, 1894) (see Kornicker, 1975:4), but slightly larger (Migs lse2a)): Size in mm: USNM 157334, length 5.0, height 4.0; USNM 157335, length 5.0, height 4.0; USNM 157336, length 5.0, height 3.9; USNM 157337, length 4.9, height 3.9; USNM 157338, length 4.9, height 3.9; USNM 157339, length 4.9, height 3.8; USNM 157340, length 4.8, height 3.9. First antenna: Ist joint: medial surface with long hairs proximally near VOLUME 90, NUMBER 4 899 middle, on ventral half all along joint, and distally on dorsal half of joint. 2nd joint: medial surface with long proximal hairs near dorsal margin and shorter hairs near ventral and terminal margins; lateral surface with long hairs near ventral margin and 9 distal bristles with short marginal spines; dorsal margin with 6 bristles (proximal 3 of these bare, or with short spines, others with long marginal spines). 3rd joint: long dorsal margin with 14 spinous bristles; short ventral margin with 1 short bristle with short mar- ginal spines. 4th joint: ventral margin with 4-5 spinous terminal bristles (inner of these small); dorsal margin with 1 long spinous bristle. Sensory bristle of 5th joint with 2 short proximal filaments and 12 or 13 longer distal filaments. 6th joint short, with few medial spines and 1 long, stout, medial bristle reaching well past a-claw of 7th joint. 7th joint: a-claw about same length as 5th joint, bare with rounded tip; b-bristle about same length as sensory bristle of 5th joint, with about 11 filaments; c-bristle reaching past tip of sensory bristle, with about 14 filaments. 8th joint: d- and e-bristles bare, about same length as sensory bristle, almost twice length of a-claw; f-bristle bent dorsally, about same length as b-bristle, with about 10 filaments; g-bristle slightly shorter than c-bristle, with about 15 filaments. Second antenna (Fig. 7b): Protopodite: medial surface, ventral and dorsal margins with long hairs (more abundant ventrally). Endopodite 3- jointed: Ist joint with 9-13 proximal bristles and 5 or 6 distal bristles; 2nd joint with 0-2 bristles; 3rd joint with 1 long terminal bristle (for distribu- tion of bristles on 7 specimens see Table 3). Exopodite: medial surface of Ist joint with minute short straight terminal bristle; minute faint spines present along ventral and dorsal margins (not observed on all limbs); all bristles with natatory hairs; bristles of joints 2-8 and 3 long bristles of 9th joint with well developed spines, much larger than those of adult male; 9th joint with 5 bristles, dorsal of these short; joints 4-8 with basal spines in- creasing in length on distal joints; basal spine of 8th joint about half length of 9th joint; lateral spine of 9th joint about same length as basal spine ot Sth joint; lateral and dorsal margins of joints 2-8 with short spines forming row (long dorsal hairs present on adult male, absent on female). Mandible: Coxale endite similar to that of adult male. Basale: endite with about 7 bristles near end (one of the bristles much longer than others), a row of 16 triaenid bristles along ventral margin, and 21 dwart bristles near dorsal margin (the most distal of these longer and stouter than others); ventral margin of basale with 13 triaenid bristles and 1 long distal bristle with long proximal and short distal spines; 2 proximal and 2 distal (almost terminal) dwarf bristles present on medial side close to ventral margin; dorsal margin of basale with 19 short bristles (distributed all along margin) and 2 long spinous terminal bristles; medial surface with long hairs distributed similarly to those of adult male, and short spines forming rows 900 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON VOLUME 90, NUMBER 4 901 near dorsal margin. Exopodite hirsute, extending just past distal end of Ist endopodial joint, with 2 ventral bristles, both with short, faint, marginal spines (distal bristle extending just past tip of exopodite, proximal bristle about twice length of distal bristle). Endopodite: 1st joint with 9 spinous ventral bristles (2 of these small); dorsal margin and medial surface (near dorsal margin) of 2nd joint with numerous bristles; ventral margin with 4 spinous subterminal bristles (bases on medial surface) and 2 long spinous terminal bristles; claws and bristles of end joint similar to those of adult male. Maxilla: Epipodite and endites similar to those of adult male. Basale: lateral side with 1 short proximal bristle near middle; 11 short bristles present on medial side just distal of, and dorsal to, bases of bristles of endite III; medial side near dorsal margin with 17-20 subequal proximal bristles; distal dorsal margin with 1 long bristle with short, faint, marginal spines, and 10 or 11 short bare bristles; ventral margin with medial row of 23-25 short bristles (proximal of these longer than distal) followed by 1 long and 2 medium length spinous bristles, and 1 long spinous terminal bristle; medial side with 7-12 short distal bristles forming row; dorsal margin of basale hirsute; hairs also present on medial surface. Exopodite consisting of minute lobe with 3 bristles (2 short, 1 longer, reaching past middle of Ist endopodial joint). Endopodite: Ist joint hirsute, with 1 short, bare, bristle near middle of anterior margin, and 1 long beta-bristle with short marginal spines; end joint with 6 bristles with short marginal spines. Fifth limb: Dorsal margin of comb slightly convex, hirsute, with 9-11 proximal small bristles. Lateral side of comb with 1 long, stout, spinous bristle anterior to 1 long, slender, spinous bristle; 3 or 4 minute bristles present between the 2 long bristles (bases of minute bristles close to base of the long slender bristle); 6 short bristles present near ventral margin ot comb in vicinity of base of long slender bristle; 2 short spinous bristles present near ventral margin at anteroventral corner of comb. Ventral mar- gin of comb with 3 rows of spinous bristles (middle row longer than medial and lateral rows), except for anterior 15 bristles which form single row. < Soo Fig. 3. Cycloleberis christiei, furca. a—c, Paratype, USNM 157336, adult female: a, Left lamella and main claws of right lamella; b, c, Medial view of part of right lamella and lateral view of part of left lamella showing concavities between main claws and position of Ist secondary claw. d, e, Holotype, USNM 157334, ovigerous female, lateral views of parts of left and right lamellae. f, g, Paratype, USNM 157318, adult male, medial view of part of left lamella and lateral view of part of right lamella. h, i, Para- type, USNM 157292, adult male, medial view of part of right lamella and lateral view of part of left lamella. j, k, Adult male from station LB 447Q, specimen lost, lateral view of part of right lamella and medial view of part of left lamella. WASHINGTON VOLUME 90, NUMBER 4 903 Sixth limb (Fig. 2e): 1-3 bristles in place of epipodial appendage, re- maining bristles of limb not counted, but limb appearing similar to that of adult male. Seventh limb: Each limb with about 117 bristles, 53-64 bristles on each side; each bristle with up to 8 bells; most rings with 2 bristles, 1 on each side, but many distal rings with 4 bristles, 2 on each side. Terminus with opposing combs, each with about 28 pectinate bristles of several types. Furca (Fig. 3a—e): Number and distribution of claws similar to those of adult male; depth of concavity in lamella between main claws 1 and 2, 30-55% of greatest width; depth of concavity between claws 2 and 3, 59- 93% of greatest width. Rod-shaped organ (Fig. 2b): Elongate, broadening posterior to suture (suture proximal to middle) and then tapering to rounded tip. Eyes: Lateral eye pigmented, well developed with about 70 ommatidia (5 rows, each with about 14 ommatidia), about % length of lateral eye of adult male (Fig. 2c). Medial eye pigmented (Fig. 2b), about same size as lateral eye, and about same size as medial eye of adult male. Posterior of body and upper lip (Fig. 2d): Similar to those of adult male. Description of adult male.—Carapace similar to that of adult male of Cycloleberis squamiger (Scott, 1894) (see Kornicker and Caraion, 1974:47), but slightly larger (Figs. 4, 5a). Size: USNM 157318, length 5.5 mm, height 3.8 mm; USNM 157319, length 5.5 mm, height 3.8 mm; USNM 157290, 2 adult males, length 5.3 mm, height 3.8 mm, length 5.3 mm, height 3.8 mm. First antenna (Figs. 5b, 6a): 1st joint with medial and lateral hairs near ventral margin (not shown in Fig. 6a). 2nd joint: dorsal margin with 4 or 5 bare bristles; lateral surface with 8 bare bristles; medial surface with long proximal hairs near dorsal margin. 3rd joint: long dorsal margin with 13 spinous bristles (not all bristles shown on illustrated limb); short ventral margin with | short bare bristle. 4th joint: ventral margin with 5 spinous terminal bristles (inner of these very small); dorsal margin with 1 long, spinous, terminal bristle. Sensory bristle of small 5th joint with abundant filaments (about 9 terminal filaments stouter than others). Medial bristle of small 6th joint bare, about equal in length to combined lengths of joints < mers Fig. 4. Cycloleberis christiei, paratype, USNM 157319, adult male, outside views of right valve: a, complete valve, length 5.5 mm; b, Rostrum and incisur, « LOQ; ec, Poste- rior end of valve showing bristles forming vertical row, fossae containing small bristle, and short curved ridges below fossae, <100; d, Detail of bases of bristles forming vertical row, from c, X1,000; e, Detail of fossae and ridges, from b, 650; f, Detail from b, * 2,000; g, h, Pores bearing bristles on anteroventral part of valve, g, \ 2,000, h, 3,000. Micrographs reduced to 55% for publication. Fig. 5. Cycloleberis christiei, paratype, USNM 157318, adult male: a, Complete specimen, length 5.5 mm; b, Proximal filament on f-bristle of lst antenna; c, Medial view of coxale endite of left mandible; d, Lateral view of comb of right 5th limb; e, Detail of dorsal margin in d; f, Detail of bristles near ventral margin in d; g, Lateral view of process and 2 bristles on dorsal margin of comb of left 5th limb, anterior to left; h, Medial eye and rod-shaped organ; i, Outline of left lateral eye; j, Anterior end of right lobe of upper lip. VOLUME 90, NUMBER 4 905 P . i S Vitex. ai ~ ' ‘4 niet y, LF WD Zip ne a —_———— LW: ae \ ——— — f x. U Fig. 6. Cycloleberis christiei, adult male. a, Lateral view of right Ist antenna of specimen from station LB 447Q, specimen lost. b-—-c, Paratype, USNM 157292: b, Lateral view of process on dorsal margin of left 5th limb; c, 7th limb. d, Paratype, USNM 157318, distal part of rod-shaped organ. 4-8. 7th joint: a-claw bare, with base on small pedestal; b-bristle about 3% times length of a-claw, with about 15 short marginal filaments (extreme tip missing on illustrated specimen); c-bristle very long, tip missing on specimen examined, remaining part with about 40 short filaments. Sth joint: d- and e-bristles well developed, longer than a-claw (1 of these with tip missing on illustrated appendage); f-bristle very long, extreme tip miss- ing On specimen examined, remaining part with about 46 short filaments 906 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON a eS SS SR [pS FP as ean Sa eee ae Ree SSS oe Fig. 7. Cycloleberis christiei. a—b, Developmental stages of endopodite of female 2nd antenna: a, Paratype, USNM 157341, A-1 instar; b, Holotype, USNM 157334, ovigerous female. c—f, Developmental stages of endopodite of male 2nd antenna, all paratypes: c, USNM 157327, A-3 instar; d, USNM 157295, A-2 instar; e, USNM 157293, A-1 instar; £, USNM 157318, adult. (proximal filaments with narrow trasparent flange); g-bristle about 6 times length of a-claw, with 8 closely spaced proximal filaments followed by 16 more widely spaced filaments. Second antenna (Fig. 7f): Protopodite bare except for small distomedial bristle. Endopodite 3-jointed: Ist joint elongate, with 6-12 short proximal VOLUME 90, NUMBER 4 907 bristles and 4-6 short distal bristles along ventral margin; 2nd joint elongate with 8 or 9 short distal bristles; 3rd joint elongate, reflexed, with 1 medium proximal bristle and serrations along inner margin of distal half of joint. Exopodite: elongate Ist joint with minute medial terminal bristle and minute spines forming rows near ventral margin; joints 3-8 with basal spines of similar length on all joints; basal bristle of 8th joint about equal in length to length of 9th joint, but smaller on some specimens; lateral spine of 9th joint about same length as basal spine of 8th joint; bristles of joints 2-8 with minute spines and natatory hairs; 9th joint with 5 bristles (dorsal of these short) with natatory hairs, some of the longer bristles may have minute spines; proximal ventral margin of bristle of 5th joint with minute spines (visible only under oil immersion); dorsal terminal corner of joints 2-8 with long hairs forming row; shorter hairs forming row along middle of distal margins of lateral side of joints 2-8. Mandible: Coxale endite (Fig. 5c): ventral branch with proximal spines on ventral margin followed by spines forming 5 or 6 oblique rows (no club- formed or finger-formed spines as on ventral branch of endite of C. galatheae described by Poulsen, 1965:262); tip of branch with 3 teeth (inner tooth with blunt tip bearing minute spines); ventral margin of dorsal branch with 4 processes pointing distally followed by 7 processes pointing proximally (the last of these very small); tip of dorsal branch more pro- longed than that illustrated for C. galatheae by Poulsen (1965:261, Fig. 87g) and terminating with minute spine; posterior bristle with base just proximal to the minute terminal spine and with few short hairs; dorsal margin of dorsal branch with hairs proximal to base of posterior bristle and with few spines just distal to middle of branch; slender bristle present near base of ventral branch. Basale: endite with 2 long stout bristles and about 8 shorter bristles near distal end, a row of 8 triaenid bristles near ventral margin, and 14 dwarf bristles along dorsal margin (the most distal of these longer and stouter that others); ventral margin of basale with 9 triaenid bristles and more distally 2 spinous bristles (the proximal of the spinous bristles only slightly longer than the triaenid bristles, and with short mar- ginal spines; the distal of the spinous bristles, long, stout, with long proxi- mal and short distal spines); 2 dwarf bristles present in distal ventral corner; dorsal margin of basale with 14 short distal bristles and 2 long spinous terminal bristles; medial surface of basale with long hairs. Hirsute exopodite extending just past distal margin of joint, with 2 bare bristles along ventral margin (proximal of these extending only short distance past tip of distal bristle). Endopodite: Ist joint with 9 spinous ventral bristles (1 of these minute); dorsal margin and dorsal halt of medial side of 2nd joint with numerous bristles (about 65); ventral margin of 2nd joint with 6 terminal bristles; end joint with 3 long stout claws and 3 bristles (1 long, lateral, 2 short ventral). 908 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Maxilla: Epipodite pointed, bare. Protopodite only weakly divided into 3 endites: endite I with 3 long and 2 short bristles; endite II with 2 long and 1 short or medium bristle; endite III with 4 long bristles. Basale: 9 short bristles present on medial side just distal and dorsal to bases of bristles of endite III; medial side near dorsal margin with 10-13 subequal but fairly short proximal bristles; distal dorsal margin with 1 long spinous bristle and 7 or 8 short bristles; ventral margin with medial row of 18 minute bristles followed by 3 medium length bristles, and then 1 long terminal bristle; medial side with 8 or 9 short distal bristles forming row; dorsal margin of basale hirsute. Exopodite consisting of minute lobe with 3 bristles (2 short, 1 fairly long, with marginal hairs). Endopodite: Ist joint hirsute, with 1 short bare bristle near middle of anterior margin, and 1 long beta-bristle with short marginal spines; end joint with 6 bristles, some with short marginal spines. Fifth limb (Figs. 5d-g, 6b): Dorsal margin of comb with single rounded process; minute and variable structures present on anterior edge of pro- cess; 5 or 6 small bristles present on dorsal margin of comb proximal to anterior edge of dorsal process. Lateral surface of comb: 1 long, stout, spinous bristle present anterior to 1 long, slender, spinous bristle; 5 minute bristles present between the 2 long bristles; about 7 short bristles present near ventral margin of comb in vicinity of base of long slender bristle; 2 short bristles present near ventral margin at anteroventral corner of comb. Hairs absent from dorsal margin of comb except on anterior edge where numerous. Ventral margin of comb with 3 rows of spinous bristles (bristles of middle row longer than those of medial and lateral rows), except for anterior 12 or 13 bristles which form single row (bristles not shown in il- lustrations). Comb without strong internal sclerotization reported for comb of male Cycloleberis squamiger (Scott, 1894) by Kornicker and Caraion (1974:56, 57, 58, Figs. 30a, 31h). Sixth limb: Medial side of anterodorsal corner of protopodite with about 9 minute, broad spines and more slender spines forming several rows; medial side of anterior margin proximal to upper suture with spinous bristles forming 3 rows (inner row with 19 bristles, middle row with 17 bristles longer than bristles in inner row, outer row with 24 bristles more slender and slightly longer than bristles in inner row but shorter than those in middle row); medial side of anterior margin between upper and lower sutures with 4 bristles in inner row, 1 bristle in middle row, and 6 bristles in outer row; about 17 bristles forming row along anterior edge between lower suture and anteroventeral corner of skirt; ventral edge of skirt with numer- ous long and short bristles; posterior end of skirt with 5 or 6 hirsute bristles; 3 short bare bristles in place of epipodial appendage; limb hirsute. Seventh limb (Fig. 6c): Each limb with about 97 bristles, 48-49 bristles on each side; each bristle with 2-8 bells; most rings with 2 bristles, VOLUME 90, NUMBER 4 909 1 on each side; some distal rings with up to 4 bristles, 2 on each side. Ter- minus with opposing combs, each with about 25 pectinate bristles of several types. Furca (Fig. 3f-k): Each lamella with 3 stout claws followed by 8 (rarely 9) bristlelike secondary claws; Ist and last secondary claws slightly shorter and more slender than other secondary claws; lst secondary claw on left lamella close to base of 3rd main claw, on right lamella separated by space from 3rd main claw; concavity between Ist and 2nd main claws shallower than that between 2nd and 3rd claws; depth of concavity in lamella be- tween main claws | and 2, 26-44% of greatest width; depth of concavity in lamella between main claws 2 and 3, 59-68% of greatest width; main claws with medial and lateral row of teeth along posterior margin, teeth about same length; Ist and last secondary claws with closely spaced fine teeth, other secondary claws with more widely spaced coarse teeth proxi- mally and fine closely spaced teeth distally; main teeth with few hairs on or near anterior margins; medial hairs present at bases of main claws; a fringe of hairs present along edge of lamellae following secondary claws. Rod-shaped organ: Elongate, broadening posterior to suture near middle and then tapering to rounded tip (Fig. 6d). Rod-shaped organ of USNM 157318 aberrant having short distal segment with teeth at tip (Fig. 5h). Eyes: Lateral eye pigmented, well developed with about 67 ommatidia (5 rows, each with 11-14 ommatidia) (Fig. 5i). Medial eye pigmented, smaller than lateral eye (Fig. 5h). Posterior of body: Hirsute but without dorsal process. Upper lip (Fig. 5j): Each lobe hirsute but without spines. Ontogeny and variability—In order to be able to compare the ap- pendages of the new species with those of Cycloleberis galatheae Poulsen, which is known only from juveniles, ontogenetic changes and variability were studied in selected characters (number of filaments on the sensory bristle of the 1st antenna, number of bristles on the endopodial joints of the 2nd antenna, and number of distal bristles on the dorsal margin of the basale of the maxilla) of the adult male and female of C. christiei, as well as on the A-1 to A-3 instars of the male, and the A-1 instar of the female. The presence of bristles on the 7th limb of the A-3 male indicates that it is not younger than a stage IV instar (see Kornicker, 1969:3). Sensory bristle of 5th joint of Ist antenna (Table 1): The sensory bristle of the A-3 male bears 2 short proximal and 7 or § long distal filaments. The sensory bristle of the A-2 male bears 2 short proximal and 10 long distal filaments. The sensory bristle of the A-1 male bears 2-4 short proximal and 10-12 long distal filaments. The adult male bears abundant filaments of which about 9 terminal filaments are thicker than the others. The sensor bristles of the A-1 female and the adult female bear 2 short proximal fila- ments. The sensory bristles of the A-1 female bear 7-12 and of the adult 910 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Table 1. Number of filaments on sensory bristle of lst antennae and on distal dorsal margin of maxilla of females and males. (nd—no data, na—not applicable.) Distal dorsal Sensory bristle of Ist antenna ; margin of basale Proximal (short) Distal (long) of maxilla Left Right Left Right Left Right Adult females USNM_ 157334 D 2 13 nd nd 12, 157340 nd 2 nd 12 nd 12 157336 2 2 13 1 10 ‘fal A-1 females 157341 2 2 12 De nd nd 157342 nd 2 nd 7 nd 8 Adult males 157318 na na na na il 9 157292 na na na na Zl 8 A-1 males 157324 2 nd nd 10 7h nd 157294 nd 33 nd i, 8 nd 15323 3 nd 12 nd 8 nd 157293 4 nd 12 nd 7 nd A-2 male 157295 2 2 10 10 6 6 A-3 males 157326 nd nd nd nd nd 4 157330 nd 2 nd 8 nd 5 ais nd 2 nd 7 nd 4 * proximal of the distal bristles with space between its base and base of 11 remaining bristles. female bear 12 or 13 long distal filaments. The sensory bristle of the A-1 male bears more short proximal filaments than the sensory bristles of the adult and A-1 females. Endopodite of male 2nd antenna (Table 2): In instars examined younger than the A-3 instar, the male and female could not be discrimi- nated. The endopodite of the A-3 male (Fig. 7c) bears 5-8 short proximal and 2-3 short distal bristles on the Ist joint, no bristles on the 2nd joint, and 1 long subterminal bristle on the smaller 3rd joint (a minute spinelike terminal bristle was found on the 3rd joint of 1 of the 7 limbs examined). The subterminal position of the bristle on the 3rd joint is the basis for fo Ud YOLG opsiuiq | ) ILSUG FHT ytd yn rei S 0 pu \ yu «I pu I es O pu 0 pu 1 I 0 pu © pu 9 pu 0 pu 0 C pu : pu (orn ie pu pu s C ur GG 8%) SLELSI 0 pu I pu © pu | (77 ie =e ) I 0 9 GG 6G) IttLs pu 0 pu 0 pu - pu fee Ait LST pu pu pu C'S 63) OFELSI 0 pu I I pa 0) 6 pe 9 : (SZ 6S) BEE 1c pu ; pu | pu SARE oT T pu C ff pu ) ys 9'Z) SZELSI pu Bo I pu 8 pu es 36) LOELST "4 x1 pu I I pu G 66) DCLLEI P el pu as pu , pu [pres I I pu 6 pu ; sajRul £-Y ee pe LE I Y pu (8°S “G'€) ELELST «1 pu I pu pu ¢ 8 jie ee u I pu pu) SZELe pu I P I pu 7 8 ; JLE I ‘ pu t pu 7 pu if . (pu) CCELST pu be g pu By IT pu (8°S 9'€) 66CLET all pu I E Be 8 (L'G S'€) LOCLET «1 pu I pu 7 (6° “S'€) S6SLST pu BIL pu V pu C ee pu pu pu sa[eul Z-V pu I pu Vv pu OL pu (pu) PCELCI «I pu V pu & pu (9'€ “G'Y) ESELST pu off I pu PT pu 8 (pu Cae _m pu I = g pu Or pu Nee a SOD ¥ - P oa & pu cs (c'¢ ‘g'F) 96ZLST 3 0 : oF T : oe OT (SE ($"P) POCLET rs Pm om ¢ (SE ‘SF) LESLET = 0) 0 jes 9 pu Cc = 0 i I Pe Cc - Eo OI sojeul [-V Z 0 I : 8 8 ; P 8 pu (pu) Z6ZLST =) : 6 ’ 9 9 = (p¥) T6SLST = SOE ee ee OL ZI Ee ‘g'G) 6TELST a ma al LG Fo Gc) SIELST WN 5 [e311 SHER IL 3 Nsa = jeulxoig Re ele 13 Ssoyen APY. S : past] HU BRT BS EO! Oa ——_—_—_—_—_— yeasiq (WIE UT YIP “Y{BUe qurol puoses ie A Geer ee ee & suoumtoeds I) yurol 4sitq (‘eyep Ou— pu) ‘soyetr jo ovuUu eS) yue pug jo s}zurof ,erpodopus uo soaz}sit iq jo JaqunNn -* CI CLAD 912 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON | Table 3. Number of bristles on endopodial joints of 2nd antennae of females. (nd— | no data.) | First joint oe Od RED Second Third Proximal Distal joint joint Specimens Left Right Left Right Left Right Left Right Adult females USNM 157334 (5.0, 4.0) Me 13 6 6 0 0 1 1 15783on (OF 40) i, nd 6 nd 2 nd it nd 157336 (5.0, 39) 12 12 5 5 i i 1 1 Nass (GES, 89) Ie nd 5 nd 1 nd 1 nd aBSO (4594 3.9) 12 nd 6 nd 1 nd iL nd 157339 (4.9, 3.8) 13 nd 5 nd 0 nd if nd 157340 (4.8, 3.9) nd 9 nd 5 nd 0 nd 1 A-1 females WS7S4Ul (BS Be) 2 nd 5 nd i nd 1 nd IE (LOL Bil) nd 12, nd 5 nd 0 nd 1 157349 (4.0, 3.1) nd 8 nd 5 nd 0 nd I 157350 (4.0, 3.2) 9 nd 4 nd 0 nd 1 nd Tes) (CLO, 89) nd 10 nd 4 nd 0 nd 1 identifying the A-3 instar as a male, because the long bristle of the 3rd joint is terminal on the female. The endopodite of the A-2 male bears 7-11 short proximal and 3 or 4 short distal bristles on the Ist joint, no or 1 distal bristle on the 2nd joint, and 1 long proximal and 1 short spinelike bristle on the 3rd joint (Fig. 7d). The 3rd joint on the A-2 instar is much longer than that joint on the A-3 instar relative to the lst and 2nd joints. The endopodite of the A-1 male bears 6-11 short proximal and 3-5 short distal bristles on the Ist joint, 4 or 5 short distal bristles on the 2nd joint, and 1 long proximal and | minute spinelike terminal bristle on the 3rd joint (Fig. 7e). The length of the 3rd joint of the A-1 male is about 65% of the combined length of joints 1 and 2, compared to about 48% on the A-2 male. The length of the 3rd joint of the A-1 male is about 80% of the length of the long proximal bristle compared to about 38% on the A-2 male. The endopodite of the adult male bears 6-12 short proximal and 4-6 short distal bristles on the Ist joint, 8 or 9 short distal bristles on the 2nd joint, and a medium length (2 or 3 times length of short distal bristles on 2nd joint) proximal bristle on the 3rd joint (Fig. 7f). The 3rd joint is reflexed on the 2nd, its tip is pointed, and it bears serrations along the inner mar- gin. The function of the endopodite of the adult male is to clasp the fe- male during copulation. VOLUME 90, NUMBER 4 913 Endopodite of female 2nd antenna (Table 3): The morphology of the joints and the position of the bristles is similar on the endopodites of the instars and on the adult, but the number of bristles on the Ist and 2nd joints differ. The 3rd joint always bears a long terminal bristle. The endop- odite of the A-1 female bears 8-12 short proximal and 4 or 5 short distal bristles on the Ist joint and no or 1 short bristle on the 2nd joint (Fig. 7a). The adult female bears 9-13 short proximal and 5 or 6 short distal bristles on the Ist joint and no or 1-2 short bristles on the 2nd joint (Fig. 7b). Distal bristles on dorsal margin of basale of maxilla (Table 1): The basale of the A-3 male bears 4 or 5 distodorsal bristles, the A-2 males 6, the A-1 male 7 or 8, and the adult male 7 to 9. The basale of the A-1 female bears 8 distodorsal bristles and the adult female 10 to 12. Remarks concerning Cycloleberis galatheae Poulsen, 1965.—Poulsen (1965:260) described this species from 2 specimens, an A-1 female (holo- type) from False Bay, South Africa, and a juvenile male paratype (A-1 or A-2 instar) from Beira, Mozambique. The Ist endopodial joint of the 2nd antennae of the female bears 2-3 proximal bristles (Kornicker, 1975:13), whereas the Ist endopodial joint of the 2nd antennae of the juvenile male bears 10 proximal bristles. The variability of the number of bristles on the Ist endopodial joint of the 2nd antenna has been studied for Cycloleberis squamiger (Scott, 1894) by Kornicker (1975:14, Table 1) and for C. christiei, herein (Tables 2, 3). Extrapolating these data to C. galatheae we conclude that the female and male referred to that species by Poulsen are not con- specific. The juvenile male from Mozambique is referred herein to C. christiei, but only questionably, because the distal dorsal margin of the basale of the maxilla of the Beira specimen bears only 4 bristles compared to 6 on the A-2 male and 7 or 8 on the A-1 male of C. christiei. More speci- mens from the vicinity of Beira will have to be studied before the identity of the species of Cycloleberis living there can be known with certainty. Comparisons.—The comb of the 5th limb of the adult male of the new species, C. christiei, differs from that of C. squamiger in not having a rounded protuberance anterior to the middle process on the dorsal margin and, also, in not having an internal sclerotized structure. The A-1 female of C. christiei differs from the A-1 female of C. galatheae in having 8-12 proximal bristles rather than 2-3 (Kornicker, 1975:13) on the Ist endopodial joint of the 2nd antenna, and in having more than 4 distal bristles on the dorsal margin of the basale of the maxilla. The sensory bristle of the 5th limb of the Ist antenna of the A-1 female of C. christiei bears 2 short proximal filaments compared to 3 on the sensory bristle of the A-1 female of C. galatheae, but the variability of this feature is not known. The adult female of C. christiei bears 9-13 proximal bristles on the Ist endopodial joint of the 2nd antenna compared to only 1-4 bristles in this place for C. squamiger. 914 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Literature Cited Day, J. H. 1959. The biology of Langebaan Lagoon: A study of the effect of shelter — from wave action. Transactions of the Royal Society of South Africa 35(5):475— 547. Kornicker, Louis S. 1969. Morphology, ontogeny, and intraspecific variation of Spinacopia, a new genus of myodocopid ostracod (Sarsiellidae). Smithsonian Contributions to Zoology 8:1-50, 26 text-figs., 6 pls. 1975. Ivory Coast Ostracoda (Suborder Myodocopina), Smithsonian Con- tributions to Zoology 197:iii + 46 pp., text-figs. 1-32. , and Francisca Elena Caraion. 1974. West African myodocopid Ostracoda (Cylindroleberididae). Smithsonian Contributions to Zoology 179:iii + 78 pp., text-figs. 1-43. Miller, G. W. 1906. Die Ostracoden der Siboga-Expedition. Siboga-Expeditie 30: 40 pp., 9 pls. Leiden: E. J. Brill. Poulsen, Erik M. 1965. Ostracoda-Myodocopa, 2: Cypridiformes-Rutidermatidae, Sarsiellidae and Asteropidae. Dana-Report 65:484 pp., 156 text-figs. Copen- hagen, Carlsberg Foundation. Scott, T. 1894. Report on Entomostraca from the Gulf of Guinea, collected by John Rattray, B. Sc. Transactions of the Linnean Society, Zoology, Ser. 2, 6:1-161, 15 pls. Skogsberg, Tage. 1920. Studies on marine ostracods, 1: Cypridinids, halocyprids and polycopids. Zoologiska Bidrag fran Uppsala, Supplement, 1:1—784, 153 text-figs. - a a PROC. BIOL. SOC. WASH. 90(4), 1977, pp. 915-920 OCCURRENCE OF THE SUBGENUS HOLOTHURIA (HOLOTHURIA) IN THE GULF OF MEXICO (ECHINODERMATA: HOLOTHUROIDEA) David L. Pawson and Thomas C. Shirley Abstract.—Pawson, D. L., Department of Invertebrate Zoology, Smith- sonian Institution, Washington, D.C. 20560, and T. C. Shirley, Department of Zoology and Physiology, Louisiana State University, Baton Rouge, Louisiana 70803.—The nominate subgenus of Holothuria was believed to be restricted to the eastern Atlantic and Mediterranean area. Recent dis- covery of populations of Holothuria dakarensis Panning in 2 localities in the Gulf of Mexico considerably extends the known range of the subgenus. A brief description of specimens from the Gulf of Mexico is given, together with ecological notes. The holothurian genus Holothuria comprises approximately 110 species and is widely distributed in shallow tropical and subtropical seas. In a recent review of the genus, Rowe (1969) recognized 17 subgenera. Several of these subgenera are relatively widely distributed in the Indo-Pacific and Atlantic oceans. Others have a more restricted distribution. Among the latter group is Holothuria (Holothuria), which was believed to occur only in the eastern Atlantic-Mediterranean area, for the distribution of its spe- cies was known to be as follows: H. (H.) fungosa Heller and H. (H.) massaspicula Cherbonnier—Red Sea. H. (H.) tubulosa Gmelin and H. (H.) mammata Grube—Mediterranean, Canary Islands, Azores, Madeira Islands (see under “Distribution” below). H. (H.) stellati Delle Chiaje and H. (H.) helleri Marenzeller—Mediter- ranean. . H. (H.) caparti Cherbonnier—West Africa between 3°05’S and 14°095’S. H. (H.) dakarensis Panning—Senegal and Angola, West Africa. In 1972 and 1973, during an ecological investigation of Seven and One- Half Fathom Reef off North Padre Island, Texas (26°49’N, 97°19’W) one of us (T.S.) found large populations of Holothuria (H.) dakarensis Panning. Further specimens were found at Isla de Lobos, located approximately 120 km southeast of Tampico, Mexico (21°27’N, 97°13’W) in 1973 and 1976. This is the first definite record of the subgenus from the western Atlantic Ocean. In this paper a brief description of the western Atlantic specimens is given, together with some notes on their ecology. T.S. wishes to acknowledge the generosity of J. W. Tunnell, Texas A&l University at Corpus Christi and J. A. Rickner, Texas A&M University, for allowing access to their collections from Mexico. Funding for the faunal 916 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON PAD B isa) w) eo cues (a) 2 SOM 3 , fys a ose ZED 5 {See ay 4 (258) fo aS S (eB & aN SHS ail 5 Ge S a = = co) ae So A jo ae cs am a 2 oS aus MPF Cie Cle me AS IS eH = aus eB a BSS Sr a D Qc \ iN ras cy SS cae ee RO? oom =A 5 ae ‘S Qs AS A eye Cv DS = oot \ : LS Ae [Po Oo 823 be S| ae y oO pS aa Loos Ay GRY 50 um 100 um _, 50um 100 um Fig. 1. Holothuria (H.) dakarensis from Gulf of Mexico. a, Tables from body wall; b, Knobbed buttons from body wall; c, Developing perforated rods from ventral tube feet; d, Rod from tentacle; e, Rods from dorsal papillae. investigation of Seven and One-Half Fathom Reef was received from the Caesar Kleberg Wildlife Foundation and Texas A&I University at Kings- ville Faculty Research Fund. We thank Miss Ailsa M. Clark at the British Museum (Natural History) for the loan of specimens and for information on the collections in her care. Order ASPIDOCHIROTIDA Family HOLOTHURIIDAE Holothuria (Holothuria) dakarensis Panning Holothuria stellati dakarensis Panning, 1939:538, figs. 10, 11. Holothuria dakarensis—Cherbonnier, 1950:102, figs. 1-3; 1965:14. Holothuria (Holothuria) dakarensis—Rowe, 1969:153. ?Holothuria tubulosa.—Domantay, 1958:189. VOLUME 90, NUMBER 4 S17) (Domantay (1958) briefly described specimens of a holothurian from “Florida” which he referred to H. tubulosa Gmelin. As the specimen can no longer be found in the Allan Hancock Foundation, University of South- ern California, Los Angeles collections, and as so many of the identifica- tions made in that paper were incorrect, it is now impossible to determine exactly what species Domantay had.) Description—Largest specimen 140 mm total length. Body approxi- mately cylindrical, mouth and anus terminal. Dorsal surface conspicuously different from ventral, with numerous evenly scattered conical papillae 1-3 mm long. Dorsolaterally, few papillae (approximately 10 on each side) conspicuously larger than others. Ventral tube feet numerous, generally scattered, but with tendency to be aggregated into radii in bands 4-6 tube feet wide. When tube feet aggregated into bands, intervening radii con- spicuously naked. Body wall relatively thin, leathery, rough to touch due to presence of numerous ossicles. Color in life light to dark brown, ventral side paler; lateral papillae and ventral tube feet light brown to white; dorsal surface with 2 rows of irregu- larly spaced darker brown blotches; 6-8 blotches in each row (fig. 1). In alcohol specimens dark brown, feet and papillae light brown; dark dorsal blotches less conspicuous. Ossicles in body wall tables and buttons. Dorsal and ventral ossicles sim- ilar, although dorsal ossicles larger than ventral, and ventral ossicles in- clude slightly more numerous buttons. standard mean (um) range (um) deviation st. error Tables Height ventral 52 42-63 1.55 0.99 Height dorsal 63 58-67 0.63 O21 Width ventral 68 G30 Io} 0.35 Width dorsal 86 79-96 tals 0.35 Buttons Length ventral 95 83-100 1.82 0.61 Length dorsal 126 100-163 4.57 1.52 Width ventral 39 33-46 0.84 0.28 Width dorsal 47 38-48 1.64 0.55 Tables (Fig. 2a) well developed, with 4 central perforations in base sur- rounded by up to approximately 12 marginal perforations of varying size. Some tables more or less reduced, with only 1 or 2 marginal perforations. Edge of disc with numerous short sharp projections. Spire short, with 1 crossbar and with 12 projections, regularly arranged. 918 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 2. Holothuria (H.) dakarensis from Gulf of Mexico. Dorsal aspect of specimen 130 mm long. Buttons (Fig. 2b) not common anywhere in body wall, somewhat vari- able in shape, usually with 3 pairs of elongate perforations. Surface of buttons with numerous rounded knobs. Tentacles contain rods (Fig. 2d) with extremely numerous small, short spines, which have tendency to be arranged in rows. Rods greatly variable in size, with a range of approximately 50-550 um. Some rods have lateral perforations which are almost obliterated. Dorsal papillae with endplates. Walls of papillae contain typical tables and rods (Fig. 2e) with holes along 2 sides and some small surface spines. Rods variable in size, up to approximately 400 um in length. Ventral feet with endplates. Deposits similar to those of dorsal papillae. Develop- mental stages of perforated rods common (Fig. 2c). Remarks.—In most respects these specimens conform well with the current concept of the species H. (H.) dakarensis. The only consistent dif- ference lies in the height of the tables. In West African specimens, table height is in the range 60-80 um (Panning, 1939) or 70-85 um (Rowe, 1969). In the present material, height of the tables is 42-67 wm, with means of 52 ym (ventral) and 63 um (dorsal). This difference is not considered to be sufficient to warrant referral of the Gulf of Mexico specimens to a new spe- VOLUME 90, NUMBER 4 919 cies, but separation at the subspecies level might prove to be desirable once the range of variation of the species in this subgenus is better known. Rowe has already commented upon the remarkable similarities between the species dakarensis, tubulosa, mammata and stellati, and has suggested characters which can be used to distinguish them. In height of the tables, the Gulf of Mexico specimens of H. dakarensis tend to overlap with H. mammata and H. tubulosa, but they differ in other characters. Distribution.—In the Gulf of Mexico, this species has been collected at 2 localities, Seven and One-Half Fathom Reef, Texas, and at Isla de Lobos, Mexico. In the eastern Atlantic H. dakarensis is definitely known only from Senegal (Panning, 1939; Cherbonnier, 1950) and Angola (Cherbonnier, 1965) to depths of 17 m. Rowe (1969, p. 154) noted that H. dakarensis also occurs at the Azores and Canary Islands. These records were apparently based upon specimens in the collections of the British Museum (Natural History). Reexamination of these specimens (1956.5.29.46, Fayal, Azores and 1966.7.13.5, Canary Islands) revealed that their ossicles fall well within the size range for H. tubulosa or H. mammata; they are considerably smaller than those of H. dakarensis. Ecology.—Seven and One-Half Fathom Reef is a sedimentary rock struc- ture lying 3.2 km from North Padre Island, Texas, in 14 m of water. The reef has been the site of numerous faunal investigations from 1968 to 1975. More complete descriptions of the reef and its associated fauna have been given by Felder (1973), Tunnell (1973), Tunnell and Chaney (1970) and Thayer, La Rocque and Tunnell (1974). Holothuria (H.) dakarensis was most abundant where steeply angled rock surfaces interface with the surrounding sandy bottom. Other individuals were also found partly buried in sediment in crevices and on sand under rocks throughout the reef. In July 1973 the species had a 20% frequency of occurrence and an average abundance of 0.26 individuals/m? in quadrats located within 3 parallel transects that traversed the reef approximately perpendicular to its long axis. Specimens were also collected on numerous occasions (by T.S.) in 1972 and 1973 and (by others) in 1968. Isla de Lobos lies in 28 m of water and is one of the northernmost emergent coral reefs in the western Gulf of Mexico (Rigby and McIntire, 1966). Holothuria (H.) dakarensis was collected at approximately 1 m depth on a calcareous sand and coral rubble bottom around the hull of a shipwreck on 5 June 1973. Additional specimens were collected from Lobos Reef in 1976 (J. W. Tunnell, personal communication). The saxicolous habits of H. (H.) dakarensis have undoubtedly contributed to its late discovery in the western Atlantic. Most invertebrate surveys have employed trawls or dredges and thus have avoided banks and reets to prevent loss of collecting equipment. The species will probably be found in other suitable habitats in the western Atlantic. 920 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Literature Cited Cherbonnier, G. 1950. Note sur Holothuria dakarensis Panning. Bull. Mus. Hist. Nat. Paris 22(1):102-108, 3 figs. —. 1965. Holothurides. Result. Scient. Exped. Oceanogr. Belge Eaux Cot. Afr. Atlant. Sud 3(11):1-24, 11 pls. Domantay, J. S. 1958. Some holothurians from Florida in the collection of the Allan Hancock Foundation. Agra Univ. Jour. Res. (Sci.) 7(2):181—202. Felder, D. F. 1973. ame, ventral view; 26. Same, close-up. VOLUME 90, NUMBER 4 939 ! = ee Fig. 27. Rhysophora ardeoceras: 27. Fourth and fifth abdominal sterna of male, ventral view. terior setae of fourth and fifth abdominal segments conspicuously larger, with oblique-dorsal orientation. Fourth sternum of male (Fig. 27) with median, oval, membraneous area; male genitalia (Figs. 22, 23) with surstyli robust, bearing some spinules; aedeagus bilobed, left lobe bearing a spinelike, acutely-pointed process. Type-material—Holotype male, labelled: “COSTA RICA: Guanacaste Prov. 14kmNE Tilaran, 05 June 1973; Erwin & Hevel, Central American Expedition, 1973.” Allotype and 1 male paratype with same label data as holotype. Other paratypes as follows: MEXICO: 1é, Tepic, 2 July 1956, R. & R. Dreisbach; 12, Jalisco, Barranquillas, 3 February 1964, E. I. Schlinger. EL SALVADOR: 1¢6, Santa Tecla (Nueva San Salvador), 12 km NW, October, 1953, W. B. Heed. COLOMBIA: 22 2, Buenaventura, 2 November 1950, Michelbacher and Ross. SURINAM: 12, Paramaribo, February, 1968, F. D. Bennett and H. Zwolter. The holotype is deposited in the National Museum of Natural History, Smithsonian Institution, Wash- ington, D.C., type number 75360. Etymology.—Ardeoceras is of Greek derivation and is a combination of the nouns ardis, meaning “point of an arrow’ and ceras, meaning “horn, in 940 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON RHYSOPHORA ARDEOCERAS Fig. 28. Rhysophora ardeoceras: 28. Distribution map. allusion to the long, pointed antennae of specimens of this species. The name stands in apposition to the generic name. Geographic distribution (Fig. 28).—Rhysophora ardeoceras occurs throughout Middle America and into northern South America between 20° north latitude and the Equator. Natural history—Gary Hevel (personal communication) collected the specimens of the primary type-series by sweeping vegetation in a roadside ditch. Phylogenetic Considerations The cladogram (Fig. 29) and accompanying list of character evidence (Table 1) summarize the relationships between the species of Rhysophora. The monophyly of Rhysophora is confirmed by several synapomorphies (characters 1-4, the apomorphy of each, determined by ex-group com- parison). None of these were used to characterize the genus in previous treatments, nor were other character states used which are unique to the genus as characterized above. Perhaps this is why Cresson (1946) con- ditionally placed Psilopa umbrosa Loew and P. caeruleiventris Loew in Rhysophora when these two species could not be accommodated within other Psilopine genera. Now, the generic concept of Rhysophora is more firmly established and Loew’s Psilopa species have been removed (Mathis and Wirth, 1977). VOLUME 90, NUMBER 4 94] Fig. 29. Argumentation scheme for the hypothetical phylogeny of the genus R/iy- sophora. I recognize two species-groups in Rhysophora despite the small number of known species. The groups are easily recognized and the monophyly of each is reasonably established as indicated in the cladogram (characters 5-6, 11-13). The relationship of Rhysophora to other ephydrid genera (mainly Psilopini) is not well understood. I have not been able to identify character states which would indicate a sister-group relationship with another genus or group of genera. Consequently, I have included in the key below those genera that are related patristically but whose cladistic relationships have not been clarified. These genera are recognized as follows: PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 942 UOI}BJUSLIO enbi[qosiop YyIM pur 199.17] SnHOUBIQUIOUL BIB [BAO ueIpow pepunor Ajoynoe yoryq ‘sno.opoorun orydioup Ayjenxes UMOIG-YSTAB.IB WYstey oho YNOFJ-9UO Oe BUIse19Ar oyeUIsIVUla ATPBOIg pessoidun Ajdoep yU98.1dATp I9UUT SSO] 10 F[ey-9uo juosoid Ud][OMS piryq} pure puodes 0} Ae[LUIIS pezHolspos Ajeinua pepuno.r Apuny[q yor|q Jepuleutet ay} ‘asue.i0 YSIMoyad ‘oped uonsz0d [e.QUeAo.1e}sod Ie]TUIS Sojeulof pue So[eUl PIES | Jystey 9A9 Y}UVE}IJ-9UO UBY} sso] BWO}SIda JO IopUIVUAI YIM [OA] JusplAd Ft possoiduit ApAMoyTpeys Jugs I9AIp-oyeurpoo.id Jouur 0} [enba quosqre ISIV}ISVG I1OF PUR PIU 0} AR]TUITS JUOUISS [PUTWIOpge Y}ANOF JO Uls1eW LOL19}sod SuOTe UOT}VUSIAO puv OZIS [eS gjeu JO UINUIO}s [eUTWIOpe Y}ANO,T Juoulses [euUSJUe piIy} Jo odeys JuOUIseS [RUUO}JUR PTY} FO UOT}eIO[OD sie} JO UOTPBALO[OT) UOTFeLO[OO [P19U9f) JYystey [Rue UIsIvUL [eIORF [P.A}UOA BIAOF peuuo}Uuy SOPSLIq 1e][I00}s0d Fo UOLUITIC IPSLIG [VOAGA 19}NoO JO YSU] ole ul jo snsiejyiseq pult{ fo DVJINS [VIPUOAOLOJUR BUOCTR IvJOS ged ‘oyeylo JO MOL oyYI[osULIy ole ul jo snsiezyiseq puly fo edeys o1dAjode ord A} 0tsayd S9}V}S jAo}pORIeYD 1dJOBICYT) ‘plioydoshyy Jo satoods oy} Fo sisAyeue onjeyAyd ur pasn sajzejs JoJoRIeYyO puUP Sid}OR.1eY-) WAI eal VOLUME 90, NUMBER 4 943 1. Prescutellar acrostichal setae large, their insertions relatively sepa- rated, anterior of, or aligned with intra-alar setae. 2. Strong supra-alar seta present. 3. Specimens generally robust. 4. Thoracic vestiture usually dull, pollinose. This assemblage of genera is intermediate between the tribes Discocerini and Psilopini as Cresson (1942) characterized them. In an earlier paper, Mathis and Wirth (1977) suggested that Discocerini and Psilopini, sensu Cresson (1942), should be combined in the absence of discrete, identifying character states. I have since studied these genera and now believe that Discocerini is a monophyletic assemblage if the genera included in the tribe are limited to those Cresson recognized in his various synopses (1942, 1945, 1946a, 1946b). The characters which establish the monophyly of Discocerini, however, were not used by Cresson (1942). These are: 1. Prescutellar setae small, inserted close together and behind or aligned with the intra-alar setae. 2. Mesopleuron lacking small, up-curved seta near the posterodorsal corner. 3. Reclinate fronto-orbital bristle inserted slightly anterior of larger, proclinate fronto-orbital as viewed from a lateral aspect. 4. Smaller proclinate fronto-orbital setae, if present, inserted anterior to larger proclinate bristles. Genera included in Discocerini are: Discocerina Macquart, Hydrochasma Hendel, Polytrichophora Cresson, Diclasiopa WHendel, Hecamedoides Hendel, Pectinifer Cresson, and Ditrichopora Cresson. This restricted concept of Discocerini leaves Psilopini with the genera Cresson normally included in that tribe, plus genera frequently included in Discocerini, i.e. Hostis Cresson and Paratissa Coquillett. Because I have not identified synapomorphic character states for Psilopini, it could be paraphyletic. Erecting a third tribe for the intermediate genera would partially alleviate the problem because the remaining psilopine genera lack a supra-alar bristle. However, this action would still leave the status of the third tribe unresolved. Key to Rhysophora and Related Psilopine Genera with a Large Supra-alar Bristle and with Prescutellars Well Separated 1. Vein Rz,3 basad of anterior crossvein with 3-4 setae above 2 — Vein Rz,3 basad of anterior crossvein bare above { Aristal branches numbering 3-4; postocellar setae with proclinate- divergent orientation; wing hyaline or uniformly infumated; alula normal 3 — Aristal branches numbering 8-15; postocellar setae with latero- bo 944 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON clinate-divergent orientation; wing guttate, generally dark brown with white spots; alula reduced Actocetor Becker Four fronto-orbital bristles, first 2 bristles proclinate, third latero- clinate, fourth lateroreclinate; 1 pair of infrafrontal bristles anterior OO of ocellar bristles Paratissa Coquillett — Three fronto-orbital setae, first bristle proclinate, second reclinate, third proclinate; lacking intrafrontal bristles Hostis Cresson 4, Wing immaculate, hyaline or uniformly infumated; aristal branches numbering 8-14; thoracic coloration mostly black, subshiny to shiny; outer vertical bristle with length half or less that of outer vertical bristle; hind basitarsus of male swollen, bearing a row of fine, cilia- like, pale setae along anteroventral surface; postocellar setae latero- proclinate Rhysophora Cresson — Wing with anterior margin darkened, usually brownish black or wing guttate, dark, brownish black with white spots; aristal branches numbering 4-6; thoracic coloration mostly dull, grayish- pollinose; hind basitarsus of male similar to mid and fore basitarsi, lacking row of cilialike setae; postocellar seta with proclinate-di- vergent orientation ) 5. Wing guttate, dark, brownish black with white spots; costal vein index greater than 1:0.80 Guttipsilopa Wirth — Wing darkened along anterior half, posterior half hyaline; costal vein index less than 1:0.60 Nesopsilopa Mathis and Wirth Acknowledgments I am grateful to the following curators and their respective institutions for the loan of specimens: Dr. Pedro W. Wygodzinsky, American Museum of Natural History (AMNH); Mr. Guy E. Shewell, Canadian National Col- lection (CNC); Dr. Howard V. Weems, Jr., Florida State Collection of Arthropods (FSCA). For their critical review and constructive commentary of this paper, I thank Drs. Wayne E. Clark, Richard C. Froeschner, and Willis W. Wirth. Mr. George C. Steyskal advised me on the formation of the specific epithets. Technical assistance was ably provided by Miss Mary Jacque Mann, scanning electron micrographs; by Mr. Victor E. Krantz, wing photos; by Miss Hollis B. Willams, arrangement of locality data and maps: by Ms. Debbie E. Townsend, illustrations 1-2, 13-14, and 20-21; and by Miss Anne Halpem, for carefully typing the final draft of this paper. Literature Cited Cresson, E. T., Jr. 1924. Descriptions of new genera and species of the dipterous family Ephydridae. Paper VI. Entomol. News 35:159—-164. ———. 1942. Synopses of North American Ephydridae (Diptera). I. The Subfamily Psilopinae, with descriptions of new species. Trans. Amer. Entomol. Soc. 68: 101-128. VOLUME 90, NUMBER 4 945 1945. WASH. 90(4), 1977, pp. 946-962 DEONTOSTOMA COPTOCHILUS N. SP., A MARINE NEMATODE (LEPTOSOMATIDAE) FROM THE FOOT CAVITY OF THE DEEP-SEA ANEMONE ACTINAUGE LONGICORNIS (VERRILL, 1882) W. Duane Hope Abstract—Hope, W. D., Department of Invertebrate Zoology, National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560.—Deontostoma coptochilus, a new species of marine nematode (Leptosomatidae), is described. The specimens upon which the description is based were extracted from sediment within the foot cavity of the deep- sea anemone Actinauge longicornis (Verrill, 1882) collected at 1,000 m depth off the coast of Uruguay. This is a new depth record for a species of this genus. The associations between various species of Deontostoma and other marine organisms is discussed. Numerous specimens of a new species of Deontostoma have been dis- covered in the foot cavity of the deep-sea anemone, Actinauge longicornis (Verrill, 1882), collected during cruises of the German research vessel Walter Herwig. The sea anemone was the subject of an earlier investigation by Dr. K. Riemann-Ziirneck (1973) and the nematodes were made ayail- able for the present study through the courtesy of Dr. Franz Riemann of the Institut fiir Meeresforschung in Bremerhaven, West Germany. Materials and Methods All specimens were fixed in 6-8% formalin, then preserved in 80% ethanol. Subsequently, each specimen was processed by the slow method into anhydrous glycerine and mounted on glass slides for preliminary study. Ten males (NMNH Nos. 53542 thru 53551), and 11 females (NMNH Nos. 53567 thru 53577), including the holotype and allotype, were selected for the measurements listed in Table 1. Two males (NMNH Nos. 93542 and 53564) have been decapitated for the purpose of preparing face views for light microscopy, and their spicula and gubernacula, as well as the ventromedian supplement of NMNH No. 53564, were excised and perma- nently mounted for photomicrography. The right spiculum of a third male (NMNH No. 53565) was also excised and mounted on a slide, while the head was prepared for SEM. The heads and tails of 2 additional males (NMNH Nos. 53558 and 53566) were also prepared for SEM. The procedure for SEM preparation has been described elsewhere (Hope, 1974). VOLUME 90, NUMBER 4 947 Fig. 1. Head and neck regions of the holotype (A) and allotype (B) of Deontostoma coptochilus in lateral view. Scale applies to A and B and equals 20 um. Deontostoma coptochilus, new species Figs. 1-8 Specimens studied.—25¢ ¢, 372 2; 18 00 Description.—Body small for Deontostoma (Table 1), gradually tapered anteriorly and posteriorly from mid-body region; head bluntly rounded (Figs. 1 & 2). Tail conical with blunt terminus (Figs. 4 & 5). Cuticle smooth. Head with 6 labial papillae and 10 cephalic setae. Short lateral, subdorsal, and subventral cervical setae between cephalic suture and region of ocelli. Setae posterior to nerve ring, in 1 subdorsal, 2 sublateral and 1 sub- ventral row on each side of body. Cephalic capsule (Fig. 1) without prominent anterior lobes or tropis and with usual 6 posterior lobes. Each posterior lobe broadened at its posterior end with or without small anteriorly directed tines. Posterior end of each lobe rounded to truncate with but few shallow notches or incisions. Intralobar lacunae absent. Anterior margin of cephalic capsule with or 948 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 2. Head region of Deontostoma coptochilus in lateral view. A, Photomicrograph showing the dorsal odontium (LOD) of the left subventral mandibular ridge and dorsal onchium (ON). Scale is 20 um; B, SEM picture showing odontia on dorsal (DOD) and right (ROD) and left (LOD) subventral mandibular ridges. Scale is 10 um. VOLUME 90, NUMBER 4 949 Yj UH), "yy Yjjpy Yy Yj, Wifi Fig. 3. SEM face view of Deontostoma coptochilus showing microlabia (ML) man- dibular ridges (MR) and dorsal (DOD) and left subventral (LOD) odontia. The dorsal onchium is barely visible in the stoma. Scale is 5um. without adjacent band of intracuticular reticulation. Amphid circular to oval. Stoma (Fig. 3) triangular with orifice enclosed by 1 dorsal and 2 sub- ventral microlabia. Dorsal microlabium nearly divided by mandibular ridge bearing paired odontia; each subventral microlabium completely divided by mandibular ridge; each subventral mandibular ridge bearing paired odontia. All 6 odontia anteriorly directed and bluntly conical. In single specimen measured, interval between tips of dorsal odontia 5.7 jum; and 6.7 and 6.9 wm between tips of each member of right and left subventral pairs respectively. Dorsal wall of stoma with single large onchium (Figs. 1A, 1B and 2A); tip of onchium at level of cephalic setae. Esophagus conical and with paired ocelli; each ocellus comprised of spherical lenslike structure and reddish-brown pigment spot. Dorsal esoph- ageal gland orifice approximately midway between posterior margin of cephalic capsule and ocelli (Table 1). Tail terminus with caudal gland pore. Caudal glands outstretched and extending anterior to rectum (Fig. 4A). Males.—As in general description. Subventral supplements (Fig. 5B) in 950 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Table 1. Morphometric data for Deontostoma coptochilus. Character a b c Total length (mm) Dist. to amphid* Amphid length Amphid width Head width Cephalic capsule- length* Cephalic setae Distance to DEGO (absolute)+ Distance to DEGO (% of dist. to ocellus) Distance to ocellus* Length of esophagus (mm)* Body width at base esophagus Midbody width Distance to vulva (absolute; mm)_ Distance to vulva (% of body length) Ova length Ova width Spicula length (arc) Gubernaculum length Anal vent-vmsi Number of preanal setae-right Number of preanal setae-left Preanal setae-length Mammate svs-right Mammate syvs-left Cloacal body diameter Anal body diameter Tail length All absolute measurements in ym unless indicated otherwise. * Measured from anterior margin of head. N oO ac Os Os OS O35 Os OS Oy O38 Os Os OS OS Os O35 Os OS Be BB RP Bo Re pe ke +0 +O +0 40 40 +0 40 +0 +0 +0 +0 +0 40 140 40 +0 Os Os Os OS Re Re +O +0 +0 +0 go ce OY eG Ce YO Num- ber of vari- ates 21 21 20 21 21 21 21 21 21 44 20 10 9 64 10 10 10 10 21 Range 36.9-58.1 6.00-8 .32 85.6-166.1 9.59-13.07 12.7—20.9 9.58-18.2 8.75-17.2 35.4—48.0 29.0-44.5 5.0-10.0 44.5-80.0 39.0-68.1 98.0-132.7 1.42-1.93 185-226 215.5-273.9 5.28-7.16 48.2-60.0 198.60-363.00 102.70-—308.00 175.5-193.8 110.6—132.6 81.6-104.9 15-24 16-25 11-23 2-4 3-4 118.0-142.8 100-116 75.4-118.0 Mean 46.06 6.88 115.17 129 als WTA 12.25 41.86 34.63 dol 61.83 54.59 113.39 1.66 205.2 246.7 6.20 54.43 284.63 173.64 188.81 120.07 96.55 19.4 19.88 TAO 3.20 3.60 131.19 106.20 WaT: Stan- dard devi- ation 5.43 0.49 20.00 0.96 2.56 2.45 2.61 3.07 4.18 1.41 8.94 6.76 9.10 Oars Wu lL 1726 0.60 3.56 53.68 68.52 6.42 6.43 6.93 2.87 3.28 3.37 0.60 0.48 7.70 3.00 12.08 } Distance from anterior margin of head to dorsal esophageal gland orifice. + Distance from anal vent to ventromedian supplement. Stan- dard error 1.18 0.10 4.47 0.21 0.56 0.53 0.57 0.67 Oo 0.21 1-99 1.51 1.98 0.02 2.4 3.8 0.18 1.07 14.88 19.00 2.03 2.03 2.19 0.90 1.09 0.42 0.18 0.15 2.43 1.56 2.70 951 VOLUME 90, NUMBER 4 so Ss eT <<, . S SS with photo- View ‘ral late in ontostoma coptochilus graphy (B). of De micrography (A) and scannin equals 30 um. tails Female Fig. 4. applies to A and B and > ral Sc TO ’ ctronmic g ele PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON VOLUME 90, NUMBER 4 953 one longitudinal series on each side of body extending anteriorly from just posterior to cloaca. Each series comprised of setae typically located on top of low transverse folds of cuticle. Setae at anterior end of series interspersed with papillae on top of prominent, mammate evaginations of cuticle and underlying hypodermis. Posteriormost 3 or 4 evaginations very prominent; anterior to this, evaginations distinctly smaller. Cuticle of evaginations without closely packed, internal rodlike structures as occur in Deontostoma californicum Steiner & Albin, 1933. Ventromedian supplement (Figs. 5A, B; 6A, B; 7A—C) a raised oval disc a short distance (Table 1) anterior from cloacal vent. Disc bears depres- sion slightly posterior from its center with ridge extending anteriorly. Floor of depression raised, to form stout, flat-topped peduncle with central pore. Disc circumscribed by oval field of minute striae on surface of cuticle (Fig. 6B) and flanked anteriorly and posteriorly by zones of intra- cuticular reticulations; anterior zone approximately circular in shape, pos- terior zone lenticular (Fig. 6B, C). Spicula (Fig. 8A) scimitar-shaped with angular bend near distal end; calomus short in relation to lamina. Lamina comprised of 3 ribs; ventral rib wide between calomus and distal third of lamina; dorsal lamina widest distal to angular bend. Calomus with lateral, anteroventrally directed pro- cess. Capitulum not enlarged or otherwise set off from calomus. Guber- naculum (Fig. 8B) duplicate; corpus of each half consisting of parallel bars fused at each end. Proximal end broad for muscle attachment; distal end with several distinct transverse striations. Corpus without a crus. Male reproductive system diorchic; testes opposed. Females.—As in general description. Reproductive system amphidelphic and reflexed. Types.—Holotype (Male)—NMNH No. 53543. Allotype (Female) NMNH No. 53575. Paratypes: Males—NMNH Nos. 53542 and 53544 thru 53566. Females—NMNH Nos. 53567 thru 53574 and 53576 thru 53601. Juveniles—NMNH Nos. 53602 thru 53619. Type-habitat—Sediment contained within the foot cavity of the sea anemone Actinauge longicornis (Verrill, 1882). Type-locality—The benthos at 1,000 m depth off the coast of Uruguay near the mouth of the Rio de la Plata, 34°28’S, 51°40’W. Distribution —The localities from which D. coptochilus have been col- < Fig. 5. Posterior ends of male Deontostoma coptochilus. A, Photomicrograph showing spiculum (SP), gubernaculum (G) and ventromedian supplement (VMS). Seale is 80 um; B, Scanning electronmicrograph showing setiform subventral supplements (S), ventromedian supplement (VMS), and mammate evaginations bearing papillae (P). Scale is 50 um. 954 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON VOLUME 90, NUMBER 4 95 Ul Fig. 7. Ventromedian supplement of Deontostoma coptochilus. A, Photomicrograph showing the peduncle (P) and ridge (R); B, Photomicrograph of the intra-cuticular reticu- lations anterior (AR) and posterior (PR) to the ventromedian supplement; C, Drawing of ventromedian supplement and reticulations. In all 3 cases anterior is uppermost. Scale applies to A—C and equals 5 um. lected, as well as the depth and numbers of individuals for each station are given in Table 2. Etymology.—A combination of the Greek copto, meaning cut, and cheilos meaning lip, and referring to the division of each lip into 2 parts by the mandibular ridge. Relationships.——The cephalic capsule in the genus Deontostoma is too similar between species to be a reliable diagnostic character when used alone. It is, however, an easy structure to observe and figure and, as a re- sult, in some species it is the only structure for which useful data exists. Because of this, it currently provides one of the better known characters for recognizing groups of species within the genus, although these are not necessarily groups of common phylogenetic ancestry. With this consideration in mind, D. coptochilus may be likened to the < Fig. 6. Ventromedian supplement of Deontostoma coptochilus. A, SEM showing its position relative to the setiform subventral supplements, and its orientation. The posterior end of the animal is beyond the upper right. Seale is 30 «am; B, Details of the ventromedian supplement showing the peduncle (P) with its central pore, ridge (R and field of surface striae (S). Scale is 5 «um. 956 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON a Fig. 8. Copulatory apparatus of Deontostoma coptochilus. A, Photomicrograph of left spiculum showing the lateral process (arrow) on the calomus; B, Photomicrograph of the left half of the gubernaculum showing the striae (arrow) at its distal end, and the absence of a crus. Scale applies to A and B and equals 40 um. following species in which the cephalic capsules have slightly rounded to truncate posterior lobes and/or no intralobar lacunae: D. anchorilobatum (Allgén, 1947), D. demani (Mawson, 1956), D. elongatum (Ditlevsen, 1926), D. hopei Coles, 1977, D. karachiense (Timm, 1959), D. magnificum (Timm, 1951), and D. papillatum (Linstow, 1903). While D. anchorilobatum is not known to have intralobular lacunae, it can be distinguished from D. coptochilus by the very rounded shape of the posterior lobes. Little else is known of this species since it was described from females and juveniles and the types apparently no longer exist. D. demani differs in that the dorsal lip bears “about six” small odontia, whereas it bears only 2 odontia in D. coptochilus. The lobes of the cephalic capsule of D. elongatum resemble those of D. coptochilus more closely than any of the above species. Un- fortunately, data is incomplete regarding the structure of the stoma and male copulatory organs. However, the total body length (20 mm), distance be- tween the anterior margin of the head and the eyespot (190 um), length of the spicula (230 um), distance between the cloacal vent and ventromedian supplement (216 um) are substantially different from those of D. coptochilus (Table 1). D. karachiense is dissimilar by the striae at the distal end of the gubernaculum being on a thick lateral ridge, and by the presence of a well developed crus. D. magnificum differs by the presence of a well de- veloped crus, the dorsal odontia are nearer the dorsolateral corners of the | VOLUME 90, NUMBER 4 95 —~l Table 2. Collection data for Deontostoma coptochilus and the number of individuals of each sex and juveniles (6) from each station. Walter Herwig Number of station numbers Coordinates Depth (m) specimens 422, 35°40’S, 52°40’W 270 20S Zao Vie 431 34°28’S, 51°40’W 1,000 ag) LOO 16 68-1 34°36’S, 51°58’W 300 264 229 61lée 68-2 34°36’S, 51°58’W 300 13g 82 6684 34-60 35°43’S, 52°43’W 1,000 12 586 mouth, and the mandibular ridges do not divide the microlabia. While little is known about the anatomy of D. papillatum, the length of males and females is given as 23.30 mm in the original description, and Filipjev (1916), in his study of the type-specimens, gave the length as 22.63 and 22.78 mm. Finally, D. hopei differs by the apparent absence of an onchium, rounder tail terminus, presence of a well developed crus, and its greater length (¢ = 22.50 mm; 22 = 19.60-26.10 mm). All other species of Deontostoma have prominent locules in the posterior lobes of the cephalic capsule. Biological associations——There are many accounts in the literature of free-living marine nematodes being associated in some way with marine organisms, either plants or animals. Most of the well documented cases are from ecological studies of the relationship between nematodes and the thalli of algae or blades of sea plants (Colman, 1940; Hopper & Meyers, 1966, 1967a, 1967b; Ott, 1967: and Wieser, 1951, 1952, 1954 and 1959), and/or between the nematodes and the rhizoid with its trapped sediment (Moore, 1971; Wieser, 1959). These studies have shown positive correlations between the algae and their associated nematodes and that each of certain kinds of nematodes may be peculiar to specific regions of the same algae, i.e. the tips of the thallus, the stems, or the rhizoid, ete. Species of Deontostoma have not been found in any of the aforementioned ecological studies, although members of the closely related genus Thora- costoma have. On the other hand, species of Deontostoma have been re- ported in numerous faunal studies as associates of a diversity of organisms (Table 3). It is not known if there is a specific ecological relationship be- tween any species of Deontostoma and its respective associate. However, the facts that species of this genus do occur in free sediment, and that at least some species, particularly D. angustifissulatum and D. antarcticum, have been found in association with such a diversity of organisms (assum- ing accurate identification of the nematodes), suggests that if an ecological association occurs, it is facultative at best. PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 958 EGET “LOSOlAA ESGT “TOSOLAA EGET “19SerA\ LOGI “BAOUOze] I LOGI “eAOUO Ze] ZIGT ‘Aoffoars 6S6T “UosTTV 6S61 “USsiTV SCG “UOsMe|\ SCE “UOSMeL\ S61 “SisU] GC6I “UOsMEY, Y UOAOYY}IIS CCB ‘UOSMPIY Y UAOYYOIS PO6GT “URWed SGGT “UosMes QG6T “uosmeyy GGG ‘UOSMBIY XY UWOAOTYYOS QC6T “UOsSMeYy QG6I “UOsMe\y QGBI “UOsMe|\ ef “‘phouy op eviqeg ‘ovoryD jeueg a[tyD ‘seoojyions) seysyT aD “seoojytons) svysy] pues, sjyousle7, 1eyeo1d pueys— Aoudy 1907e0.15) ysvoyn ueULInN|\ spurs, puepyey spurlsy puepye] pues, otrenboryy purys] ugjonsiey ‘O1y.p ouuro[ puey wey. purysT ugjonsioy “Avg ueYyIq.oyy purysy] ugjensioy “Avg uPYyiqloy as, Attopuopuo'y ‘UILIOT, FO Avg puvysyT pareopy “OAOD) seiyVy pues] ugjons.loy ‘o1y,p ouuvo[ purys] ugpnsioy “Aeg uvylqiloy MZSGoP9 S:SEL9 WEOL9 “S.9FL9 pues] plvofy ‘OAOD sepyy ul § wm 1-0 Wm 27S 9UOZ [e10}I'T QUOZ [P10}I'T OuOZ [PLOT] d1OYSe POYse AA ul QT yoroq uO uw PO-ET ul (YC {YpoRvoq Ler @) JIOYSE PoYSeE MA qYoroq uO ut (g ut JT¢G-O1¢G ut €9T DIOYSE POYSsSeAA OeS[e YIM souojs [[eUIs UO sosuods oepl[yApP pue oesye YIM souojs wor djoy oeplHAW, pure ovs[e YM sou0zs DULIDYIODS DIUDULULD'T YAK pozeLoosse pues sno} fur snoOnT YI poyetoosse pues WNIUDULINUL WUNUDYy,OYWT UT, sploziyt djoy » ovs[y UOHRULLOF sIshsolon py OvB[e UIdI8 2 SpooMvos BuOWYy OSV sasuods SUSHIOLID JW S}SVFP[OY ViduUD'T spodo.jses 0} poyorze ovrsye suowy OvB/V ovs[e SUOWY JseFP[OY srpshoosov py SyseFpfOY wnimiupy,oYyWT UPIPIOSE FO SOT. oeBV WNOVYOLD WUNOVYOMD JUD WNID)]JOIOUD WNIDINSSLUSNDUD dUdLOJOY uot BOOT yidoq o]yPLOOSSYV soloods ‘9SBO YORO OF USAID dIv DIUAoFoL puv UOTROOT “YWdop sz ‘soyeroosse ay} JO ouleu oY], ‘SUISIURSIO OULIBUL 1OYJO YP UOHRIOOSse UL puNOF Us0q AVY YY} DULOJSOJUOAGT JO soroeds oy} JO SI WY 'E aTqry, 959 Wg96T “Aydanyy LOGI ‘edozy] PLOL ‘odoxy] OL8T “uoLeyy (unoiflUusdU *G = pboifiovd *f) eCGgB[ ‘Aydin OIT6[ “19UT9}5 6S6T “uur y, CC6I “UOSMeTY X UDAOY YI} S96T Huosseyy pure [[o19 6S61 “UssT[y CSET “LOSorAA CSET “JOSolM CSET “LOST AA CSET “TOSol MA d0UdLOFOY VOLUME 90, NUMBER 4 UOJSUTYSe AA ‘RON 9p ueNf Fo Wes BIULOFIeD) “OAOAS) Olfloed Y yowog uojyjiq ROHOIRJUY “purysy ssoy aouRL] “UOpe.1qUO;y UOSIIQ “YOOY YorzsArypy BIS FTAA 0} 9ouRIJUY Je VIS s}UsIeg UR STYe IAA ‘Tyoviey “puvysy erouryy purys] Ugfonsioy ‘Aeg uePYIqIos\ BIUIOFITeD “Yyovog UoOTIC PIBIOIL) YINOS seole oruoJUuY UeS-osteredye A a[tqD ‘seore oonery 9p OJ[OF)-ouenyeoye J, gtyD ‘enbmby jo yyos NETO) ‘e[nsulueag oquimboy U01}VI0'T QUOZ [PLOT] QUOZ [V10}}I'] wl JC OUOZ [P1O}WT] JUOZ [el0WT QUOZ [R10] wg QUOZ [R10] [epasequy [242] 9p MOT MOTI Wd (OST [Peppy [epasequy yidoq peq snpyAyy wos, JUSLUIpas SYSPFP[OYW VIAALAY DIYOADJUD DIYOIMDIUD DIASOY sisdohig UIA syooL UGC Speq [essnul Japun JUsUIIpag dPB[V eRs[e pue syooy S}seFp[OYy sushIoLId py S}SPJP[OY YDpISIP DIUDUNLUDT pue DFIAav] VIFAIDY sproziyt djay plo SSeFplOy Dyjlaing Sn}}ep MOY ovs[e UseIS pony oes[e UMOIG SIG JO s}seJplOH oes[e UMOI BIG JO s}seyploy a} B1lD0ssy ——— eee as UDUOTAULY pM ULTLLOTY JAFULUAT] ASUIDUO PALJUOUL WNIUIUTZDU WnNjDqO} ISUALYIDIDY yaqaduws WNIWUAOJIIDI avIpuD]yono 5 ee satoads 5 ‘ponurjuo7 te GAL 960 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON There are reasons why specimens of Deontostoma species might tend to occur in higher numerical density in sediments associated with sessile or semisessile organisms, if indeed such is the case. First, the host organism may, in a high energy, littoral environment, provide a stable habitat by trapping or holding sediment and detritus beneath it. Second, the host organism, be it plant or animal, and the various organisms it shelters, may secrete or excrete various organic compounds that serve as substrates upon which bacteria and other microorganisms may exist. These bacteria may, in turn, be the foundation of an entire food web, of which Deontostoma is a primary and/or secondary consumer. Third, it is possible that the host provides shelter from currents or predators. Protection from currents may be especially important in the littoral zone where many of the associations are known to occur. Nothing is known about the natural enemies of Deontostoma aside from the report of juvenile specimens of D. antarcticum having been recovered from the gut of a bottom feeding fish belonging to a species of Zanchlorynchus (Mawson, 1958). Indeed, the large size of the body of individuals of Deontostoma would make them more visible prey for fish than the smaller nematodes, should they emerge, even in part, from the sediment. The apparent association between D. coptochilus n. sp. and Actinauge longicornis is unique in that it is the first report of a Deontostoma species in sediment from the foot cavity of a sea anemone, although, in view of the diversity of apparent associations in which specimens of Deontostoma are known to engage, this new account is not surprising. This is also the first report of such an association from the depths of a continental slope, the previous greatest depth having been 440 m (Vitiello, 1975). Literature Cited Allgén, C. A. 1959. Freeliving marine nematodes. Further Zoological Results of — the Swedish Antarctic Expedition 1901-1903, 5(2):1-293. | Colman, J. 1940. On the fauna inhabiting intertidal seaweeds. Journal of the | Marine Biological Association 24:129-183. | Croll, N. A., and A. R. Maggenti. 1968. A peripheral nervous system in Nematoda with a discussion of its functional and phylogenetic significance. Proceedings of | the Helminthological Society of Washington 35(1):108-115. | Filipjev, I. N. 1916. Freeliving nematodes in the collection of the Zoological Museum of the Imperial Academy of Sciences in Petrograd (In Russian with English | diagnoses). Akademiya Nauk, Zoologicheskii Muzei, Ezhigodnik, 21:59-116. | English translation by M. M. Haque, Amerind Publishing Co., New Delhi, | 1973:;1—55. { Hope, W. D. 1967. Free-living marine nematodes of the genera Pseudocella Filipjev, | 1927, Thoracostoma Marion, 1870 and Deontostoma Filipjev, 1916 (Nematoda: | Leptosomatidae) from the West Coast of North America. Transactions of the | American Microscopical Society 86(3):307—334. ——. 1974. Deontostoma timmerchioi n. sp., a new marine nematode (Lepto- somatidae) from Antarctica, with a note on the structure and possible func- | VOLUME 90, NUMBER 4 961 tion of the ventromedian supplement. Transactions of the American Microscop- ical Society 93(3):314—324. Hopper, Bruce E., and S. P. Meyers. 1966. Observations on the bionomics of the marine nematode, Metoncholaimus sp. Nature 209(5026):899-900. 1967a. Foliicolous marine nematodes on turtle grass Thalassia testudinum Konig, in Biscayne Bay, Florida. Bulletin of Marine Science 17(2):471-517. 1967b. Population studies on benthic nematodes within a subtropical sea- grass community. Marine Biology 1(2):85—96. Inglis, W. G. 1958. A new species of the nematode genus Thoracostoma from the Antarctic. Annals and Magazine of Natural History, Series 13, 1(1):45-48. Man, J. G. de. 1904. Nématodes libres (Expédition Antarctique Belge). Résultats du Voyage du S.Y. Belgica: 1-51. Marion, A. F. 1870. Recherches zoologiques et anatomiques sur des nématoides non parasites, marins. Annales des Sciences Naturelles, 13(14):1—100. Mawson, P. M. 1956. Freeliving nematodes. Section I: Enoploidea from Antarctic stations. British-Australian-New Zealand Antarctic Research Expedition 1929-— 19315 6(3):37—74. 1958. Freeliving nematodes. Section III: Enoploidea from Subantarctic stations. British-Australian-New Zealand Antarctic Research Expedition 1929— 1931, 6(14):307-358. Moore, P. G. 1971. The nematode fauna associated with holdfasts of kelp (Laminaria hyperborea) in northeast Britain. Journal of the Marine Biological Association of the United Kingdom 51:589-604. Murphy, D. G. 1965a. Thoracostoma pacifica n. sp. from the coast of Oregon. Proceedings of the Helminthological Society of Washington 32(1):106—109. 1965b. Thoracostoma washingtonensis, n. sp. eine Meeresnematode aus dem pazifischen Kistenbereich vor Washington. Abhandlungen und Verhandlungen des Naturwissenschaftlichen Vereins in Hamburg 9:211-216. Ott, J. 1967. Vertikalverteilung von Nematoden in Bestanden nordadriatischer Sar- gassaceen. Helgolander Wissenschaftliche Meeresuntersuchungen 15:412—428. Platonova, T. A. 1967. Free-living marine nematodes of the family Leptosomatidae from the European Arctic. Zoologicheskiy Zhurnal 46(6):828—838. Riemann-Zurneck, K. 1973. Actinaria des Siidwestatlantik I. Hormathiidae. Helgo- lander Wissenschaftliche Meeresuntersuchungen 25:273-325. Saveljev, S. 1912. Zur Kenntnis der freilebenden Nematoden des Kolafjords und des Relictensee Mogilnoje. Societe Impériale des Naturalists St. Petersbourg (Zoologie) 43(1):108-126. Steiner, G. 1916. Freilebende Nematoden aus der Barentssee. Zoologische Jahrbicher 39(5/6):511-676. Schuurmans Stekhoven, J. H., and P. M. Mawson. 1955. On some free-living marine nematodes from Kerguelen Island. Journal of Helminthology 29(1 2):S7—LO4. Timm, R. W. 1951. A new species of marine nematode, Thoracostoma magnificum, with a note on possible “pigment cell” nuclei of the ocelli. Journal of the Washington Academy of Sciences 41(10):331—333. Wieser, W. 1951. Untersuchungen itiber die algenbewohnende Mikrofauna marine: Hartbéden. I. Zur Oekologie und Systematik der Nematodentauna von Plymouth Osterreichische Zoologische Zeitschrift 3(3,/4): 425-480. 1952. Investigations on the microfauna inhabiting seaweeds on rocky coasts IV. Studies on the vertical distribution of the fauna inhabiting seaweeds below the Plymouth Laboratory. Journal of the Marine Biological Association of the United Kingdom 31:145-—174. 962 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON ———. 1953. The free-living marine nematodes. I. Enoploidea. Lunds Universitets Arsskrift (Avdelning 2) 49(6): 1-155. ———. 1954. Untersuchungen iiber die algenbewohnende Mikrofauna mariner Hartboden. Hydrobiologia 6(1/2):144—217. 1959. Free-living marine nematodes. IV. General Part. Lunds Universitets Arsskrift (Avdelning 2) 55(5):1-111. Vitiello, P. 1975. Deontostoma parantarcticum n. sp. et Thoracostoma ancorarium n. sp., nouvelles espéces de Leptosomatidae, (Nematoda) d’Afrique du Sud. Trans- actions of the Royal Society of South Africa 41(4):339-350. PROC. BIOL. SOC. WASH. 90(4), 1977, pp. 963-967 RANGE EXTENSIONS OF FOUR SPECIES OF CRANGONID SHRIMPS FROM CALIFORNIA AND BAJA CALIFORNIA, WITH A KEY TO THE GENERA (NATANTIA: CRANGONIDAE) Mary K. Wicksten Abstract.—Wicksten, M. K., Allan Hancock Foundation, University of Southern California, University Park, Los Angeles, California 90007.— Southern range extensions are given for Crangon spinosissima, C. styli- rostris, and Argis californiensis. Pontophilus occidentalis is reported for the first time in California. Female A. californiensis differ in size and propor- tions from male specimens reported by previous writers. The flattened dactyls of the pereiopods of Argis probably are used in digging, not swim- ming. A key to the genera of the Crangonidae in California is presented. Members of the family Crangonidae, the sand shrimps, are abundant on bottoms of sand, mud or shell along the coasts of California and Baja Cali- fornia. During identification and cataloguing of specimens of these shrimp in the collections of the Allan Hancock Foundation, extensions of the ranges of four species were found. Of the four, one has not been reported from Californian waters before, two have not been taken previously off Baja California, and one was collected south of its previous limit in California. To assist in identification, a key to the genera of the Crangonidae in this area is included. The specimens collected by the R.V. Velero IV during 1975 and 1976 were taken as part of the Southern California Baseline Studies and Analysis (FY 1975-1976) funded by the Bureau of Land Management (U.S. Depart- ment of Interior), contract number 08550-CT5-52. Pontophilus occidentalis Faxon 1893 Pontophilus occidentalis Faxon, 1893:200. Previous records—Between Mariato Point and Cocos Island, 2,169- 3,087 m; off Cocos Island, 97-1,970 m; Gulf of Panama, 103-3,382 m; off Galera Point, 2,904-3,212 m; and between the Galapagos Islands and Aca- pulco, 2,511-4,121 m (Faxon, 1895); off Chile, 1,910-2,120 m, and off Peru, 1,850-2,100 m (Zarenkov, 1976). Material. —32°54’N, 118°55’W to 33°01’N, 119°05’W; 20.3 mi, 258°T to Castle Rock Light, San Clemente Island, California, 1,754-1,825 m, lv July 1963, Isaacs-Kidd midwater trawl, mud bottom, R.V. Velero I\_ station 8791. 964 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON The single specimen obtained at station 8791 probably was taken when the trawl accidentally hit the bottom. The total length (orbit to tip of tel- son) was 32 mm. Crangon spinosissima Rathbun 1902 Crangon spinosissima Rathbun, 1902:891. Crago spinosissima: Schmitt, 1921:100, fig. 69. Previous records.—Off California and Oregon, 28-177 m, south to Point Fermin (Schmitt, 1921); off Pismo Beach, San Luis Obispo Bay, 148-203 m (Goodwin, 1952, as Crago spinosissima); Orange County, 137 m, Dana Point, 91 m (Word, 1976). Material examined.—30°31/N, 116°10’W: 3.75 mi NW of Isla San Martin, Baja California, 78 m, sand, 3 March, 1949, R.V. Velero IV station 1694, one specimen, total length 25 mm. This is the first record of C. spinosissima from Mexican waters. Crangon stylirostris Holmes 1900 Crangon stylirostris Holmes, 1900:174-175, figs. 33-35. Crago stylirostris: Schmitt, 1921: 90, fig. 61. Previous records.—Santa Cruz, California, to Chirikof, Alaska, to 48 m (Schmitt, 1921). Material examined.—35°09/N, 120°44’W: San Luis Obispo Bay, Cali- fornia, 15-26 m, rock, 4 August 1938, R.V. Velero III station 885-38, one specimen, total length 20 mm. This is a southern range extension of 1°41’ in latitude. Argis californiensis (Rathbun 1902) Fig. I Nectocrangon californiensis Rathbun, 1902:892.—Schmitt, 1921:102-103, ie a/ale Previous records.—Santa Cruz and Santa Catalina Island, 109-286 m (Schmitt, 1921); Santa Monica Bay, 183 m (Word, 1976). Material examined—CALIFORNIA, USA: 34°04’N, 119°17”W: 5.67 miles from Port Hueneme, 212-249 m, 27 April 1976, R.V. Velero IV station 24855, 3 specimens, total lengths 54, 56, and 58 mm. 33°51/N, 119°54’W: 5.67 miles to East Point, Santa Rosa Island, 172-218 m, 13 November 1975, R.V. Velero IV station 23283, 4 specimens, total lengths 50, 52, 54, and 62 mm, 2 of them ovigerous. 33°50’N, 119°53’W: 6.99 miles to East Point, Santa Rosa Island, 185-231 m, rocky bottom, 22 April 1976, R.V. Velero IV station 24798, 1 specimen, total length 37 mm. 33°41’N, 118°08’W: 3 miles VOLUME 90, NUMBER 4 965 off Seal Beach, 20 m, sandy bottom, 7 July 1940, R.V. Velero III station 1158-40, 1 specimen, total length 20 mm. 33°34’N, 119°00’W: 5% miles N of Santa Barbara Island, 140-161 m, gray sand, 29 May 1939, R.V. Velero III station 981-39, 1 specimen, broken. 33°18’N, 119°22’W: off San Nicolas Island, 179-192 m, 12 April 1940, R.V. Velero III station 1125-40, 1 specimen, total length 40 mm. 33°00’N, 118°33’W: off Wilson Cove, San Clemente Island, 111-157 m, sand and shell, 18 February 1939, R.V. Velero [II station 911-39, 1 specimen, total length 25 mm. 32°39N, 119°16’W: 43% miles to China Point, San Clemente Island, 249-259 m, “soft bottom,” 10 April 1976, R.V. Velero IV station 24684, 1 specimen, total length 37 mm. 32°39’N, 119°12’W: 40 miles to China Point, San Clemente Island, 203-240 m, rock bottom, 10 April 1976, R.V. Velero IV station 24675, 1 specimen, total length 37 mm. 32°34’N, 117°22’W: 9 miles off San Diego, 144-150 m, sand and pebbles, 23 February 1941, R.V. Velero III station 1240-41, 1 specimen, total length 34 mm. BAJA CALIFORNIA, MEXICO: 31°07’N, 116°33’W: 40.5 miles to Punta Banda, 194-222 m, 19 February 1966, R.V. Velero IV station 10990, 1 speci- men, total length 48 mm. Ten specimens taken during 1939-1941 by the R.V. Velero III off Santa Catalina Island, California also were examined. Remarks.—All of the specimens of Argis californiensis were taken in benthic trawls or dredges. Taken in the same samples were brachyurans and anomurans. The hermit crab Parapagurodes laurentae McLaughlin and Haig, 1973 was collected with A. californiensis at stations 911, 1125, and 1240, as well as off Santa Catalina Island (stations 1028, 1149, and 1150). The female specimens collected during operations of the R.V. Velero LV differ from the male illustrated by Rathbun (in Schmitt, 1921). The fe- male carapace is more inflated and the eye tubercle more elevated (Fig. 1). The females also are larger; an ovigerous female from station 23253 mea- sures 19.3 mm along the carapace instead of 8.6 mm and has a total length of 62 mm instead of 31 mm. All the specimens of A. californiensis were taken in benthic samples. Specimens of A. levior, A. ovifer, and A. dentata examined by me also were taken in benthic samples. These occurrences cast doubt on the statement that the fourth and fifth pereiopods of Argis are “natatorial” (Schmitt, 1921; Holthuis, 1955). Zarenkov (1965) noted that the tubercular conceptacle of the eyes and the flattened and broadened dactyls of the last pereiopods are adaptations for burrowing into the bottom, during which the eyes would remain above the surface. The present records of A. californiensts strongly support Zarenkov’s idea that these species have a fossorial form ol life. Neushul, Clarke, and Brown (1967) termed A. californiensis (as N. cal 966 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 1. Argis californiensis, female, life size. Velero station 23283. forniensis) an “insular endemic species.” The records from off Port Hueneme, Seal Beach, San Diego, and Punta Banda as well as Word's records (1976) from Santa Monica Bay and Orange County show that the species is not limited to the Californian islands. However, if the species is fossorial, its range may be limited by the grain size of the substrate. At § of the 12 stations at which substrate was noted, the animals were collected on sand, rock or a mixture of sand, rocks, and shell, but not mud. Because many areas of the mainland coast have muddy or silty bottoms, the species may be unable to survive there. Key to Genera of Crangonidae in Californian Waters 1. Second pereiopods lacking; rostrum elevated, with 4 spines—Para- crangon [1 species, Paracrangon echinatus Dana, 1852] — Second pereiopods present; rostrum, if present, short and without spines 2 2. Dactyli of fourth and fifth pereiopods flat and broad; eyes elevated —Argis [2 species, Argis levior (Rathbun, 1902) and Argis californiensis (Rathbun, 1902) ] -— Dactyli of fourth and fifth pereiopods not flat and broad; eyes not elevated 3 3. Second pereiopods subequal in length to other pereiopods; eyes with obvious pigment—Crangon [17 species] — Second pereiopods much shorter than other pereiopods; eyes nearly without pigment—Pontophilus [One species, Pontophilus occidentalis Faxon, 1893] Literature Cited 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 Calli- VOLUME 90, NUMBER 4 967 fornia .. . by the U.S. Fish Commission steamer “Albatross,” during 1891... . VI. Preliminary descriptions of new species of Crustacea. Bull. Mus. Comp. Zool., Harvard Univ. 24(7):149-220. — —. 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, 67 pls. Goodwin, D. G. 1952. Crustacea collected during the 1950 bottom-fish investigations of the M.V. N.B. Scofield. Calif. Fish and Game 38(2):163-181, 11 figs. Holmes, S. J. 1900. Synopsis of Califomia stalk-eyed Crustacea. Occ. Pap. Calif. Acad. Sci. VII:262 pp., 4 pls., 81 figs. Holthuis, L. B. 1955. The recent genera of the caridean and stenopodidean shrimps (class Crustacea, order Decapoda, supersection Natantia) with keys for their determination. Zool. Verhandelingen No. 26:157 pp., 105 figs. Neushul, M., W. D. Clarke, and D. W. Brown. 1967. Subtidal plant and animal communities of the southern California islands. Pp. 37-55, figs. 1-31 in Phil- brick, R. N., (ed.) Proceedings of the Symposium on the Biology of the California Islands. Santa Barbara, Santa Barbara Botanic Garden. Rathbun, M. J. 1902. Descriptions of new decapod crustaceans from the west coast of North America. Proc. U.S. Nat. Mus. 24 (1272):885-905. Schmitt, W. L. 1921. The marine decapod Crustacea of California. Univ. Calif. Publ. Zool. 23:1—470, pls. 1-50, 165 figs. Word, J. Q. 1976. Invertebrates of southern California coastal waters. Il. Natantia. El Segundo: So. Calif. Coastal Water Res. Project. 238 pp., numerous figs. Zarenkov, N. A. 1965. Revizija rodov Crangon Fabricius i Sclerocrangon G. O. Sars (Decapoda, Crustacea). Zool Zhurnal 44(12):1761-1775, 9 figs. 1976. K faune desjatinogikh rakoobraznykh priamerikanskikh vod juzhnugo polusharija. Biologija Morja 5:8-19. 6 figs. PROC. BIOL. SOC. WASH. 90(4), 1977, pp. 968-974 CAECIDOTEA CAROLINENSIS, N. SP., THE FIRSE SUBTERRANEAN WATER SLATER FROM NORTH CAROLINA (CRUSTACEA: ISOPODA: ASELLIDAE) Julian J. Lewis! and Thomas E. Bowman Abstract.—Lewis, Julian J., Department of Biological Sciences, Old Do- minion University, Norfolk, Virginia 23508, and Bowman, Thomas E., De- partment of Invertebrate Zoology, Smithsonian Institution, Washington, D.C. 20560.—Caecidotea carolinensis is described from a tectonic cave in the Piedmont Province of North Carolina. It is closest morphologically and geographically to C. richardsonae and C. catachaetus, inhabitants of the Appalachian Valley and Ridge Province. Only one asellid species has been authentically reported from North Carolina. Caecidotea forbesi (Williams), widespread in the east-central United States and in southern Ontario, was recorded from Chapel Hill, Durham Co., by Williams (1970). C. attenuata (Richardson) occurs in the Virginia part of the Dismal Swamp and may be presumed to occur in the North Carolina part also (Williams, 1970). C. forbesi and C. attenuata are both epigean species with well developed eyes, and the new species de- scribed herein is the first subterranean asellid to be reported from North Carolina. It is noteworthy that it was found in a tectonic (fissure) cave rather than in a limestone solution cave. Caecidotea carolinensis, new species Figs. 14 Material examined.—North Carolina, McDowell Co., Bennett’s Mill Cave, ca. 6 miles (9.7 km) E of Marion, leg. Cato O. Holler, 19 March 1977: Holotype, 7.4 mm é, USNM 169989, in alcohol and on 2 slides. Paratype, 3.9mm ¢, USNM 169990, in alcohol. Paratype, 5.5mm 6, USNM 169991, in alcohol and on 5 slides, exuvia in alcohol. Description of é.—Small, eyeless, unpigmented. Length up to at least 7.4 mm; body slender, linear, about 4.4 times as long as wide; coxae all visible in dorsal view. Margins of head, pereonites and telson moderately setose; shorter setae scattered on dorsum as in Fig. 1b. Head about 1.6X as wide as long; anterior margin concave; postmandibular lobes moderately produced. Telson about 1.5x as long as wide; sides subparallel; caudo- medial lobe produced as broad are. Antenna 1 reaching proximal margin of last segment of antenna 2 pe- duncle; flagellum with 9-11 segments, 3 terminal segments each bearing single esthete. Antenna 2 about 0.9X as long as body (excluding uropods) ? VOLUME 90, NUMBER 4 969 Fig. 1. Caecidotea carolinensis (d—e from holotype, other figs from 5.5 mm <¢ para ype or exuvia of same): a, Dorsal view; b, Head, dorsal (exuvia); ¢, Right antenna | exuvia); d, Antenna 2, proximal segments; e, Same, distal segments; f, Left mandible; ¥, Right mandible, incisor and spine row; h, Mandibular palp (exuvia); i, Manilla 1. yuter lobe; j, Same, inner lobe; k, Pereonite 4. 970 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON | Fig. 2. Caecidotea carolinensis (d—f from holotype, other figs. from 5.5 mm 4 paratype or exuvia of same): a, Maxilliped; b, Pereopod 1; c, Left pereopod 1, medial (exuvia); d, Pereopod 1; e, Pereopod 4; f, Pereopod 5; g, Pleopod 1. VOLUME 90, NUMBER 4 97] a Fig. 3. a-g, Caecidotea carolinensis (g from holotype, other figs. from 5.5 mm paratype or exuvia of same): a, Pleopod 2, posterior; b, Pleopod 2, tip of endopod, anterior (exuvia); c, Same, posterior; d, Pleopod 3; e, Pleopod 4; f, Pleopod 5; g, Right uropod, dorsal. h, Caecidotea richardsonae, syntype, tip of ¢ pleopod 2 endopod, anterior (from slide prepared by Steeves). last segment of peduncle about 1.5x length of preceding segment: flagel- lum with 46 (5.5 mm 4) to 55 (holotype) segments; apical segment with esthete. Mandible with 4-cuspate incisors and lacina; spine row with 11 spines in left mandible, 14 spines in right mandible; segments 2 and 3 of palp each with row of spines with robust bases and slender plumose tips. Maxilla 1, outer lobe with 13 robust apical spines and 1 subterminal seta; inner lobe with 5 apical plumose setae and 4 spinules on distal part of medial margin. Maxilliped with 5-6 retinaculae. Pereopod 1 propus 1.2-1.4x as long as wide; palm with proximal process 972 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Fig. 4. Known distribution of Caecidotea richardsonae (circles) and C. carolinensis (square). absent, its usual position occupied by 3 spines, and large blade-shaped median process separated by deep cleft from shorter, weakly bicuspid distal process; dactyl flexor margin with proximal rounded boss and distal row of spines or tubercles, depending on maturity (compare Figs. 2b and 2d). Pereopod 4 moderately setose; dactyl with 2 spines on flexor margin. Pleopod 1 longer than pleopod 2; protopod about 0.6 length of exopod, with 2 retinaculae. Exopod about 0.7 as wide as long, with about 4 setae on distal margin, 2-3 subterminal, 4 on distolateral lobe, and 4-5 on con- cave lateral margin. Pleopod 2 exopod, proximal segment with long non- plumose seta; distal segment oblong, with 9-11 non-plumose setae on margin of distal part. Endopod more than 2x as long as width at base, nearly straight; basal apophysis low, broadly rounded; tip of endopod ending in 4 processes: (1) lateral process well developed, extending disto- laterally nearly perpendicular to axis of endopod, digitiform; (2) cannula just distal to and extending obliquely across posterior surface of lateral process, about half length of lateral process, beak-shaped; (3) mesial process low, rounded, extending distolaterally posterior to cannula; (4) caudal process broadly rounded. Pleopod 3 exopod, proximal segment about 0.6 length of distal segment, with about 7 setae on lateral margin; VOLUME 90, NUMBER 4 973 distal segment with about 10 setae on straight distal margin; margins of both segments minutely serrate. Uropod rather setose; protopod about 2.6x length of exopod and 1.2% length of endopod; endopod spatulate. Female.—Unknown. Etymology.—Named after the state of North Carolina. Relationships.—Caecidotea carolinensis is closest morphologically and geographically to C. richardsonae Hay (1901), redescribed by Steeves (1963) and C. catachaetus (Fleming and Steeves, 1972). The é pleopod 2 endopod tip in all 3 species has an elongate digitiform lateral process, a beak-shaped cannula (both extending distolaterally), an inconspicuous me- sial process, and a broadly rounded caudal process. The accounts of C. richardsonae by Steeves (1963) and of C. catachaetus by Fleming and Steeves (1972) do not include a caudal process, but we found this structure to be present in slides prepared from type-material of both species. (Fig. 3h). Steeves also omitted mention of the strong sclerotization of the endo- pod proximal to the tip in C. richardsonae; this sclerotization is not found in C. carolinensis. The ¢ pleopod of C. carolinensis, C. richardsonae, and C. catachaetus are similar, all having a well developed distolateral lobe, a feature shared also by C. hobbsi, C. kendeighi, C. kenki, C. spatulata, and C. tridentata. The é pereopod 1 of C. richardsonae and C. catachaetus lack palmar processes, structures well developed in C. carolinensis. A é pereopod 1 propus sim- ilar to that of C. carolinensis is found in C. franzi (Holsinger and Steeves, 1971). Both have a rudimentary proximal process apparently situated on the posterior margin rather than the palm and represented by 3 spines, and a strong median process separated from a weakly bicuspate distal process by a deep cleft. The ¢ pleopods 1 and 2 of C. franzi, however, are radically different from those of C. carolinensis. Distribution.—C. carolinensis is known only from Bennett's Mill Cave, a small tectonic (fissure) cave located on the bank of Muddy Creek (Holler, in litt.). This cave is situated within the western Piedmont Province (Fig. 4). The localities where C. richardsonae and C. catachaetus are known to occur (Steeves, 1963; Fleming and Steeves, 1972) all lie within the Appala- chian Valley and Ridge Province, hence these 2 species are isolated from C. carolinensis by the intervening Blue Ridge Province. Since the popula- tions of C. richardsonae and C. catachaetus occur in rather disjunct areas of karst, it would not be surprising to find evidence of evolutionary di- vergence among the populations of either or both species. Acknowledgments To Dr. Cato O. Holler, who collected this interesting asellid, Dr. John E. Cooper, North Carolina State Museum of Natural History, who made 974 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON the specimens available to us, and to Dr. Brian Kensley, South African Museum, who reviewed the manuscript, we extend our sincere thanks. Facilities were kindly made available to JJL by Old Dominion University and the Smithsonian Institution. Literature Cited Fleming, Lawrence E., and Harrison R. Steeves III. 1972. Two new species of | troglobitic asellids from the United States. American Midland Naturalist 87(1): 245-249. Hay, William P. 1901. Two new subterranean crustaceans from the United States. Proceedings of the Biological Society of Washington 14:179-180. Holsinger, John R., and Harrison R. Steeves, III. 1971. A new species of subter- ranean isopod crustacean (Asellidae) from the central Appalachians, with re- marks on the distribution of other isopods of the region. Proceedings of the Biological Society of Washington 84(23):189—200. Steeves, Harrison R., III. 1963. The troglobitic asellids of the United States: the Stygius group. American Midland Naturalist 69(2):470-481. — —. 1966. Evolutionary aspects of the troglobitic asellids of the United States: the Hobbsi, Stygius and Cannulus groups. American Midland Naturalist 75(2): 392-403. Illinois. American Midland Naturalist 85(1):231-234. Williams, W. D. 1970. ‘ . PI 7 ay} Pare Ta eller P, re - r - J - “ 7 a ; —— : 1 de 4 f “ = i 4 } é J ¢ i i > ies = 4 - ) = OD : LE RSS 5 f i ) ° = i j ; ‘ - i ‘ ¥ ® ; 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, Washing- ton, D.C. 20560. 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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 84% 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 1% of a printed page. Underline singly scientific names of genera and lower categories; leave other indica- tions 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 985 - y | a Observations on specimen fixation William R. Taylor 753 Six new species and subspecies of Naesiotus from the Galapagos Islands (Pul- monata: Bulimulidae) Joseph Vagvolgyi 764 — Postembryonic development and synonymy for Eosentomon rostratum Ewing (In- secta-Protura) R. A. Durey and T. P. Copeland 778 Myodocopid Ostracoda of the Indian River complex, Florida Louis S. Kornicker 788 Buceroemersonia, a new genus of ischnoceran Mallophaga found on the hombill genus Tockus (Bucerotidae) Robert E. Elbel 798 The types and type-localities of Oreortyx pictus (Douglas) and Ortyx plumiferus Gould M. Ralph Browning 808 The status and systematic position of the species of the bopyrid isopod genus Phyllodurus Stimpson, 1857 John C. Markham 813 Ceratolana papuae, a new genus and species of mangrove-boring cirolanid isopod from Papua New Guinea Thomas E. Bowman 819 Distributional notes on Notiosorex and Megasorex in western Mexico Robert D. Fisher and Michael A. Bogan 826 On the genera Echinaster Mueller and Troschel and Othilia Gray, and the validity of Verrillaster Downey (Echinodermata: Asteroidea) Enrico Tortonese and Maureen E. Downey 829 Notes on a collection of crabs (Crustacea: Brachyura) from the east coast of Mexico Jack A. Rickner 831 Anthurids from the west coast of North America, including a new species and three new genera (Crustacea, Isopoda) George A. Schultz 839 Two new species of Ashmunella from Dona Ana County, New Mexico, with notes on the Ashmunella kochii Clapp complex (Gastropoda: Pulmonata: Poly- gyridae) Artie L. Metcalf and Richard A. Smartt 849 A new species of Synchelidium (Crustacea, Amphipoda) from sand beaches in California J. Laurens Barnard 877 Stilbomastax, a new genus of spider crab (Majidae: Tychinae) from the West Indies region, with notes on American relatives Austin B. Williams, J. Kevin Shaw, and Thomas S. Hopkins 884 Cycloleberis christiei, a new species of marine Ostracoda (Suborder Myodocopina) from Saldanha Bay and Langebaan Lagoon, South Africa Louis S. Kornicker and Rosalie F. Maddocks 894 Occurrence of the subgenus Holothuria (Holothuria) in the Gulf of Mexico (Echinodermata: Holothuroidea) David L. Pawson and Thomas C. Shirley 915 A revision of the genus Rhysophora Cresson with a key to related genera (Diptera: Ephydridae) Wayne N. Mathis 921 Deontostoma coptochilus n. sp., a marine nematode (Leptosomatidae) from the foot cavity of the deep-sea anemone Actinauge longicornis (Verrill, 1882) W. Duane Hope 946 Range extensions of four species of crangonid shrimps from California and Baja California, with a key to the genera (Natantia: Crangonidae) Mary K. Wicksten 963 Caecidotea carolinensis, n. sp., the first subterranean water slater from North Carolina (Crustacea: Isopoda: Asellidae) Julian J. Lewis and Thomas E. Bowman 968 An Atlantic helioporan coral (Coelenterata: Octocorallia) Frederick M. Bayer and Katherine Margaret Muzik 975 A new species of Sphaerodactylus (Sauria, Gekkonidae) from Isla Monito, West Indies Albert Schwartz A revision of the subspecies of Strix leptogrammica Temminck, 1831 (Aves: Strigiformes, Strigidae) S. Dillon Ripley 993 Descriptions of new taxa of crotalid snakes from China and Ceylon (Sri Lanka) Howard K. 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